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The Vascular Plants of British Columbia

Part 3 - Dicotyledons(Primulaceae through Zygophyllaceae)

and Pteridophytes

edited byGeorge W. Douglas1, Gerald B. Straley2 and Del Meidinger3

1 George Douglas 2 Gerald Straley 3 Del Meidinger6200 North Road Botanical Garden Research BranchR.R.#2 University of British Columbia B.C. Ministry of ForestsDuncan, B.C. V9L1N9 6804 S.W. Marine Drive 31 Bastion Square

Vancouver, B.C. V6T 1Z4 Victoria, B.C. V8W 3E7

August 1991

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TABLE OF CONTENTS

ACKNOWLEDGEMENTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iv

INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Format . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1Data Base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3Taxonomic Concepts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

DICOTYLEDONS (PRIMULACEAE THROUGH ZYGOPHYLLACEAE) . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

PTERIDOPHYTES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111EQUISETOPSIDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113LYCOPODOPSIDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116PTEROPSIDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121

REFERENCES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139

EXCLUDED SPECIES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154

INDEX . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159

ERRATA - PARTS 1 AND 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176

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Pteridophytes

6. Plants usually free-floating; microsporangia and megasporangia borne in separate sporocarps;leaves (in ours) less than 1 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . AZOLLACEAE (p. 125)

6. Plants rooted; microsporangia and megasporangia borne in the same sporocarp; leaves (in ours)2-15 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . MARSILEACEAE (p. 133)

5. Plants homosporous, terrestrial or epiphytic; sporangia borne on the leaves, not enclosed in sporocarps.

7. Sporangia large, borne in terminal appendages that arise from a common stalk . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . OPHIOGLOSSACEAE (p. 133)

7. Sporangia small, borne on the back or margins of the leaves or on modified fronds.

8. Fronds very delicate (resembling a leafy liverwort), blades only one cell thick; plants rhizomatouswith threadlike rhizomes, much branched and spreading . . . HYMENOPHYLLACEAE (p. 133)

8. Fronds much coarser, blades two to several cells thick; plants tufted or with rhizomes which arestouter and not threadlike.

9. Sori on or near the margin of the lower surface of the frond, covered by inrolled indusium-likemargin, or apparent margin.

10. Rhizomes coarse, much elongated, with hairs only, lacking scales; fronds scattered, verylarge and coarse, usually over 30 cm long . . . . . . . . . . . DENNSTAEDTIACEAE (p. 126)

10. Rhizomes very short, scaly, but often with hairs as well; fronds mostly clustered, muchsmaller, sometimes quite delicate, less than 30 cm long . . . . . ADIANTACEAE (p. 121)

9. Sori on the lower surface of the fronds, not covered by the leaf margins.

11. Leaves conspicuously dimorphic, fertile and sterile fronds dissimilar.

12. Sori continuous along both sides of the midrib; sterile leaves pinnate, pinnae withentire margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . BLECHNACEAE (p. 125)

12. Sori not continuous; sterile leaves pinnate-pinnatifid . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . DRYOPTERIDACEAE (p. 126)

11. Leaves not dimorphic, fertile and sterile fronds alike.

13. Indusia present, at least on young fertile fronds, sometimes only as filamentoussegments.

14. Indusia attached in the centre, margins of indusia free.

15. Rhizomes thick, short-creeping; veins not reaching the margins; indusia circu-lar or kidney-shaped . . . . . . . . . . . . . . . . . . . . DRYOPTERIDACEAE (p. 126)

15. Rhizomes slender, long-creeping; veins reaching the margins; indusia kidney-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . THELYPTERIDACEAE (p. 136)

14. Indusia attached along their margins.

16. Sori fused in a chain-like fashion along the midrib of pinnules. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . BLECHNACEAE (p. 125)

16. Sori single, rounded or oblique, not fused in a chain-like fashion.

17. Fronds once pinnate with ovate or obovate pinnae . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ASPLENIACEAE (p. 124)

17. Fronds pinnate-pinnatifid or twice (or more times) pinnate . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . DRYOPTERIDACEAE (p. 126)

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13. Indusia absent.

18. Fronds pinnatifid, the lobes not reaching the rachis . . . . . . . . . . . POLYPODIACEAE (p. 135)

18. Fronds either pinnate-pinnatifid or twice (or more times) pinnate.

19. Leaves ciliate-margined and with unicellular hairs along the rachis and costae . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . THELYPTERIDACEAE (p. 136)

19. Leaves glabrous or merely glandular, not ciliate margined . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . DRYOPTERIDACEAE (p. 126)

EQUISETOPSIDA

EQUISETACEAE

EQUISETUM

1. Shoots with cones at top of stems, rarely also at ends of the branches.

2. Fertile and sterile shoots not alike, fertile ones whitish to pinkish, unbranched or sparsely branched,fertile shoots appearing earlier than branched sterile shoots, cones appearing in spring.

3. Cone-bearing stems usually not branched, fleshy, whitish, pinkish or brown.

4. Stems 5-15 mm in diameter; stem sheaths with 20-40 teeth; cones hollow, 4-8 cm long . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. telmateia

4. Stems 3-5 mm in diameter; stem sheaths with less than 20 teeth; cones solid, less than 4 cm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. arvense

3. Cone-bearing stems green and branched, persistent, although cone and tips of stems soon wither.

5. Stem sheaths with teeth fused into 3-6 broad lobes, the number of lobes less than number of ribson stem; branches usually branched again . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. sylvaticum

5. Stem sheaths with distinct teeth, the number of teeth equal to number of ribs on stem; branchesnot branched again.

6. Stem sheaths with teeth which have a broad whitish membranaceous margin; branches fine,solid, rectangular in cross section; branch sheaths with 4 teeth . . . . . . . . . . . . . . E. pratense

6. Stem sheaths with brownish green teeth lacking whitish membranaceous margins; branchescoarse, hollow, triangular in cross section; branch sheaths with 3 teeth . . . . . . . E. arvense

2. Fertile and sterile shoots similar, fertile ones green, branched or unbranched, appearing at about thesame time as sterile shoots or later, cones appearing in summer.

7. Cones rounded at the top.

8. Stems with deep furrows; central cavity of stems less than 1⁄4 of stem diameter; stem sheaths with4-12 teeth, teeth black with conspicuous white margins . . . . . . . . . . . . . . . . . . . . . . . . E. palustre

8. Stems without deep furrows; central cavity of stems more than 1⁄2 of stem diameter; stem sheathswith 15-30 teeth, teeth with obscure, narrow white margin or early deciduous.

9. Stems firm, light green; central cavity of stems at most 2⁄3 of stem diameter; sheaths dull, theteeth of the stem sheaths deciduous leaving undulated margin of the sheath . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. laevigatum

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9. Stems softer, dark green; central cavity of stems at least 4⁄5 of the diameter; sheath shiny, thetips of the sheath with small blackish, narrowly white-margined teeth . . . . . . . . E. fluviatile

7. Cones with apiculate tips.

10. Shoots robust; stems 40-150 cm tall and about 6 mm thick, with 15-25 flat ribs, central cavitiesabout 2⁄3 of stem diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. hyemale

10. Shoots smaller; stems less than 50 cm tall and less than 4 mm thick, with less than 14 ribs, solidor central cavities up to 1⁄3 of stem diameter.

11. Stems straight or slightly curved, usually more than 15 cm long and more than 0.6 mm indiameter, hollow, central cavities about 1⁄3 of stem diameter; sheaths with at least 4 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. variegatum

11. Stems curved, contorted, usually less than 15 cm long and less than 0.6 mm in diameter,solid; sheaths with 3 or rarely 4 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. scirpoides

1. Shoots sterile, without cones.

12. Shoots branched with branches in whorls.

13. Teeth of the stem sheaths fused in 3-6 lobes; most branches branched again in a tree-like fashion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. sylvaticum

13. Teeth of the stem sheaths distinct; branches not branched again or with occasional branches.

14. Stems without ribs, central cavities about 4⁄5 of stem diameter; branches in one or very few,often incomplete, whorls . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. fluviatile

14. Stems with ribs, central cavities not more than 2⁄3 of stem diameter; branches in numerous ±regular whorls.

15. Stems thick, robust, 0.5-1.5 cm in diameter, pale green; ribs 20-40, inconspicuous;sheaths with 20-40 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. telmateia

15. Stems slender, less than 0.5 cm in diameter, green; ribs 4-18, prominent; sheaths withless than 20 teeth.

16. Branches delicate; teeth about 1⁄2 of the total length of sheaths; central cavity of stemslarger than 1⁄2 of stem diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. pratense

16. Branches coarse; teeth 1⁄4-1⁄3 of the total length of sheaths; central cavity of stems lessthan 1⁄2 of stem diameter.

17. Lowest branch internodes longer than the stem sheaths; teeth of the branchsheaths 2-4 times longer than wide, acuminate; central cavity of stems about 1⁄5of stem diameter, side cavities narrower . . . . . . . . . . . . . . . . . . . . . . . E. arvense

17. Lowest branch internodes shorter than the stem sheaths; teeth of the branchsheaths as wide as long, acute; central cavity of stems about 1⁄3 of stem diameter,side cavities almost as wide as the central cavity . . . . . . . . . . . . . . . E. palustre

12. Shoots not branched, or with irregular branches (sometimes a result of the injury of the terminal part ofthe stem).

18. Central cavity of stems about 4⁄5 of stem diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. fluviatile

18. Central cavity of stems less than 2⁄3 of stem diameter, or solid.

19. Sheaths entirely green with deciduous teeth; stems annual with smooth ridges . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. laevigatum

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19. Sheaths with a black band or entirely black or brown, teeth persistent or tardily deciduous; stemsperennial or evergreen with minutely toothed or tuberculate ridges.

20. Stems robust, more than 2.5 mm in diameter and usually over 50 cm tall; teeth tardilydeciduous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. hyemale

20. Stems finer, less than 2.5 mm in diameter and shorter than 50 cm; teeth persistent.

21. Stems straight or slightly curved, usually more than 15 cm long and more than 0.6 mm indiameter, hollow, central cavity about 1⁄3 of stem diameter; sheaths with at least four teeth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. variegatum

21. Stems curved, contorted, usually less than 15 cm long and less than 0.6 mm in diameter,solid; sheaths with three or rarely four teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . E. scirpoides

Equisetum arvense L. (E. arvense L. var. boreale [Bong.] Rupr.)Common or field horsetailHabitat/Range: Wet to mesic sandy or disturbed sites, streambanks, fields, open places and shady

forests from the lowland and steppe vegetation to alpine zones; common throughout BC; circumpolar,throughout Canada and US, MX; Eurasia, N. Africa, New Zealand.

Equisetum fluviatile L. em. Ehrh. (E. limosum L.)Swamp horsetailHabitat/Range: Shallow water at lake margins in the lowland, steppe vegetation and montane zones;

frequent throughout BC; circumpolar, N to AK, YT and NT, E to NF and S to VA, IN, IA, NE, WY, ID andOR; Eurasia.

Notes: A hybrid with E. arvense (E. x litorale Kuehl. ex Rupr.) is frequent in places where both speciesoccur.

Equisetum hyemale L. ssp. affine (Engelm.) Stone. (Hippochaete hyemalis [L.] Brubin; E. hyemale var.affine [Engelm.] A.A. Eaton, var. californicum Milde, and var. elatum [Engelm.] Morton)

Scouring-rushHabitat/Range: Moist alluvial forests, riverbanks, roadsides, and forest margins in the lowland,

steppe vegetation and montane zones; frequent throughout BC; N to AK, YT and NT, E to NFand S throughout US from FL to CA.

Equisetum laevigatum A. Br.Smooth scouring-rushHabitat/Range: Dry to mesic sandy soil, wet margins of lakes, and roadsides in the steppe

vegetation and montane zones; frequent in SC and SE BC; E to ON and S to OH, IL, OK, TX,NM, AZ, CA, and MX.

Notes: A hybrid with E. hyemale (E. x ferrissii Clute) occurs infrequently in SC BC.

Equisetum palustre L.Marsh horsetailHabitat/Range: Marshes, swamps, and streambanks and forests from the lowland and steppe

vegetation to montane zones; frequent in S BC, infrequent elsewhere; circumpolar, N to AK, YTand NT, E to NF and S to VE, CT, MI, IL, MN, ND, MT, ID and CA; Eurasia.

Equisetum pratense Ehrh.Meadow horsetailHabitat/Range: Mesic to wet river banks, open grassy slopes, bog margins, wet forests from the

montane to subalpine zones; frequent in N BC E of Coast-Cascade Mountains; circumpolar, Nto AK, YT and NT, E to MI and S to NJ, MI, IA, SD, MT and WA; Eurasia.

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Equisetum scirpoides Michx.Dwarf scouring-rushHabitat/Range: Dry to wet alpine meadows, peat bogs and dry coniferous forests in the montane to

alpine zones; frequent E of the Coast-Cascade Mountains; circumpolar, N to AK, YT and NT, Eto NF and S to CT, NY, MI, WI, IL, SD, ID and WA; Eurasia.

Equisetum sylvaticum L.Wood horsetailHabitat/Range: Moist to wet alluvial forests, meadows and swamps in the montane zone; frequent

in BC E of the Coast-Cascade Mountains; circumpolar, N to AK, YT and NT, E to NF and S toMA, PA, WV, OH, MI, WI, SD, WY, MT and WA; Eurasia.

Equisetum telmateia Ehrh. ssp. braunii (Milde) Hauke (E. maximum Lam.)Giant horsetailHabitat/Range: Moist alluvial forests, loamy banks, cliffs, and roadsides in the lowland and mon-

tane zones; frequent in coastal SW BC and Queen Charlotte Islands, rare E of the Coast-Cascade Mountains; S to CA, disjunct in MI where now possibly extirpated.

Equisetum variegatum Schleicher ex Weber & MohrNorthern scouring-rushHabitat/Range: Wet lake shores, streambanks, ditches and meadows from the lowland and steppe

vegetation to subalpine and (rarely) alpine zones; frequent in coastal (ssp. alaskanum) andinterior (ssp. variegatum) BC; circumpolar, N to AK, E to NF and S to NH, CT, PA, MI, MN, WY,CO, UT, ID and OR; Eurasia.

Notes: Two subspecies can be distinguished in BC.1. Sheath teeth brown with wide white margins . . . . . . . . . . . . . . . . . . . . . . . . . . ssp. variegatum1. Sheath teeth completely black or at most narrowly white-margined . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ssp. alaskanum (A.A. Eaton) Hult.A hybrid with E. hyemale (E. x trachyodon A. Braun) was collected on the Queen CharlotteIslands (Calder & Taylor 1968). A hybrid with E. laevigatum (E. x nelsonii [A.A. Eaton] Schaffn.)has not been reported from BC, but can be expected.

LYCOPODOPSIDA

ISOETACEAE

ISOETES

1. Corms 3-lobed; plants terrestrial, growing in grassy ephemeral pools, wet in winter and dry insummer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. nuttallii

1. Corms 2-lobed; plants aquatic, growing on bottom of lakes or on lake shores sometimes exposed insummer due to the fluctuation of water.

2. Megaspores spiny (magnification 10x or more).

3. Megaspores greatly varying in size, often aborted, surfaces resembling a brain coral; spinesblunt and dense . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. truncata

3. Megaspores uniform in size; spines sharp, not crowded together.

4. Plants flaccid; megaspore spines elongated, with pointed tips, uniform in size; micro-spores smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. echinospora

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4. Plants r igid; megaspore spines stubby and blunt, noticeably shorter and denser along theridges than on the free surface of the spore; microspores spiny . . . . . . . . . . . I. maritima

2. Megaspores smooth or with r idges, not spiny (magnification 10x or more).

5. Lowermost leaves essentially two-ranked; megaspores more than 0.5 mm in diameter;microspores rugose, more than 35 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. occidentalis

5. Lowermost leaves spirally arranged; megaspores less than 0.5 mm in diameter; microsporesspiny, less than 30 µm long.

6. Base of the leaves blackened, hyaline wing-margins of the leaves extending 1-5 cmabove the sporangium; sporangia brown spotted . . . . . . . . . . . . . . . . . . . . . . . . . . I. howellii

6. Base of the leaves green, hyaline wing-margins of the leaves not extending more than 1cm above the sporangium; sporangia not pigmented . . . . . . . . . . . . . . . . . . . . . I. bolanderi

Isoetes bolanderi Engelm.Bolander’s quillwortHabitat/Range: Shallow lakes and muddy sites in the subalpine zone; rare in SE BC, known only

from Akamina Pass; E to AB and S to NM, AZ and CA.

Isoetes echinospora Dur. (I. muricata Dur., I. braunii Dur.)Bristle-like quillwortHabitat/Range: Lakes in the lowland to subalpine zones; infrequent throughout BC; circumpolar,

N to AK, YT and NT, E to NF and S to NJ, MI, WI, MT, UT and CA; Eurasia.

Isoetes howellii Engelm.Howell’s quillwortHabitat/Range: Lake margins exposed in summer in the steppe vegetation and montane zones;

rare in SC BC, known from Kamloops Lake, Shuswap and Mara Lakes; S to MT and CA,disjunct in N UT.

Isoetes maritima Underw. (I. beringensis Komarov, I. macounii A.A. Eaton, I. muricata Dur. ssp.maritima [Underw.] Hult.)

Coastal quillwortHabitat/Range: Lake margins and shallow water in the lowland and montane zones; frequent in

coastal BC, rare in SC BC; amphiberingian, N to AK and S to WA; E Asia (Bering Island).

Isoetes nuttallii A. Br.Nuttall’s quillwortHabitat/Range: Vernal pools and ephemeral winter seepages in lowland zone; rare on SE

Vancouver Island; S to CA.

Isoetes occidentalis Henderson (I. paupercula [Engelm.] A.A. Eaton, I. lacustr is auct. non L., I.lacustr is var. paupercula Engelm., I. piperi A.A. Eaton, I. flettii [A.A. Eaton] Pfeiffer)

Western quillwortHabitat/Range: Lakes in the lowland, steppe vegetation and montane zones; frequent in S BC; S

to CO, UT and CA.

Isoetes truncata (A.A. Eaton) CluteSlashed quillwortHabitat/Range: Lake margins exposed in summer in the lowland zone; infrequent on SE Vancou-

ver Island; N to AK and S to WA.Notes: Probably a hybrid between I. echinospora and I. maritima.

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LYCOPODIACEAE

1. Sporangia in the axils of ordinary green leaves, not segregated into definite cones . . . . . . . . . Huperzia

1. Sporangia segregated into definite cones.

2. Plants long creeping with more than one fertile upright branch; sporophylls (bracts of the spike)yellowish and scale-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lycopodium

2. Plants short creeping with a single upright branch; sporophylls (bracts of the spike) green, leaf-like. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lycopodiella

HUPERZIA28

1. Leaves 0.5 mm wide, lanceolate-subulate, thin, dark green with recurved tips . . . . . . . . . H. miyoshiana

1. Leaves 1-1.5 mm wide, linear or lanceolate, firm, light or yellowish green, with straight or incurved tips.

2. Plants green, up to 20 cm tall; leaves loose, not appressed to the stem; sporangia scattered all over thestem; gemmae inconspicuous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. occidentalis

2. Plants yellowish or yellowish green, not more than 12 cm tall; leaves appressed to the stem; sporangiain the upper part of the stem; gemmae conspicuous, especially towards the top of the stem . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. haleakalae

Huperzia haleakalae (Brackenr.) Beitel (H. selago [L.] Bernh. ex Schrank & Mart. sens. lat., H. selago ssp.arctica [Tolm.] Love & Love, Lycopodium selago L. ssp. patens [Beauv.] Calder & Taylor)

Haleakala fir clubmossHabitat/Range: Dry rocky places in the alpine zone; frequent throughout BC; N to AK, YT and NT.

Huperzia miyoshiana (Makino) Ching (H. chinensis [Christ] Czer., H. selago [L.] Bernh. ex Schrank & Mart.ssp. chinensis [Christ] Love & Love, Lycopodium selago L. ssp. miyoshianum [Makino] Calder &Taylor)

Miyoshi-no fir clubmossHabitat/Range: Mesic to wet rocky places, forests, and coastal bogs from the lowland to alpine zones;

frequent in coastal BC; amphiberingian, N to AK and S to WA; E Asia.

Huperzia occidentalis (Clute) Beitel (Lycopodium lucidulum Michx. var. occidentale [Clute] L.R. Wilson)Western fir clubmossHabitat/Range: Mesic open coniferous forests, rocky places and bogs from the lowland to subalpine

zones; frequent in BC N to 56°; N to AK, E to W AB and S to ID and OR.

LYCOPODIELLA

Lycopodiella inundata (L.) Holub (Lycopodium inundatum L.)Bog club-mossHabitat/Range: Wet peat bogs and lake margins in the lowland and montane zones; frequent in coastal

BC, infrequent in C and SE BC; circumpolar (but widely disjunct), N to SE AK, E to NF, and S to MD,VA, MT, and CA; Europe, Japan.

LYCOPODIUM

1. Leaves in 6 to 10 ranks, usually more than 4 mm long.

28 Contributed by J. Beitel and A. Ceska.

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Lycopodiaceae

2. Stems upright, branched in a tree-like fashion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. dendroideum

2. Stems decumbent, or ascending, but not branched in a tree-like fashion.

3. Cones stalked, one or several on each fertile branch; midrib of the leaf extends into a long whitishhair (no hairs in var. integerrinum), the apical part of the leaves entire . . . . . . . . . . . . . L. clavatum

3. Cones sessile, one on each fertile branch; leaves without whitish hairs, the apical part of the leavesslightly serrate, or entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. annotinum

1. Leaves in 4 or 5 ranks, less than 4 mm long.

4. Stems horizontal, below ground, strongly flattened and winged; leaves on ventral side noticeablynarrower than leaves on margins or on dorsal sides of the stem . . . . . . . . . . . . . . . . . L. complanatum

4. Stems horizontal, mainly above ground, round or slightly flattened, not winged; leaves on ventral sidesimilar in size to other leaves.

5. Cones sessile; vegetative branches flattened; ventral leaves trowel-shaped; plants usuallybluish-green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. alpinum

5. Cones stalked; vegetative branches round; ventral leaves indistinguishable from otherleaves of the stem; plants usually light green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. sitchense

Lycopodium alpinum L. (Diphasiastrum alpinum [L.] Holub)Alpine clubmossHabitat/Range: Open dry to mesic areas in heath communities in the alpine zone; common

throughout BC, absent on the Queen Charlotte Islands; circumpolar, N to AK, YT, and NT, E toAB and S to MT, ID, and WA, disjunct to MI, PQ, NB, and NF; Eurasia.

Lycopodium annotinum L. (L. dubium Zoega, L. pungens La Pylaie ex Iljin)Stiff clubmossHabitat/Range: Dry to moist coniferous forests, wet peat bogs and rock outcrops in the lowland to

subalpine zones; frequent throughout BC; circumpolar, N to AK, YT, NT, E to NF and S to NC,WV, ID, and OR, disjunct in NM; Eurasia.

Notes: A variable species. In this treatment it includes L. dubium which is treated by someauthors as a separate species.

Lycopodium clavatum L. (L. lagopus [Laest.] Zinserl. ex Kuzen. = var. monostachyon)Running clubmossHabitat/Range: Dry to mesic open sites, forest clearings, forests, and bogs from the lowland and

steppe vegetation to montane zones; usually on nutr ient poor, acid soils, frequent throughoutBC, infrequent in SC BC; cosmopolitan, N to AK, YT and NT, E to NF and S to NC, MI, WI, MT,ID and N CA; Europe, Asia, Afr ica, S America.

Notes: A variable species, divided by some authors into several varieties, three of which occur inBC.1. Plants with only one cone on each fertile branch . . . . var. monostachyon Hook. & Grev.1. Plants with two or more cones on each fertile branch.

2. Tips of the leaves with a whitish hair . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . var. clavatum2. Tips of the leaves lacking a whitish hair . . . . . . . . . . . . . . . . . . var. integerrimum Spring

Lycopodium complanatum L. (Diphasiastrum complanatum [L.] Holub)Ground-cedarHabitat/Range: Dry to mesic coniferous forests and peat bog margins in the lowland, steppe

vegetation and montane zones; frequent throughout BC, rare on Vancouver Island and theQueen Charlotte Islands; circumpolar, N to AK, YT and NT, E to NF and S to NY, MI, MT, and WA; Eurasia,S America.

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Lycopodiaceae/Selaginellaceae

Lycopodium dendroideum Michx. (L. obscurum L. var. dendroideum [Michx.] D.C. Eaton)Ground-pineHabitat/Range: Dry to mesic coniferous forests, peat bogs, and rocky or sandy sites in the lowland to

montane zones; frequent on N coast and in C BC, infrequent elsewhere; amphiberingian, N to AK, YTand NT, E to NF and S to VA, IA, MT, and WA; E Asia.

Lycopodium sitchense Rupr. (L. sabinifolium Willd. ssp. sitchense [Rupr.] Calder & Taylor, Diphasiastrumsitchense [Rupr.] Holub)

Alaska clubmossHabitat/Range: Dry to mesic heath communities in the subalpine and alpine zones; common along

the coast, SW and SC BC; amphiberingian, N to AK and YT, E to NF and S to ME, NH, MT, IDand OR; E Asia.

SELAGINELLACEAE

SELAGINELLA

1. Cones stalked, circular in cross-section; leaves flat without a grooved midvein, with 2 to 4 small teethon each side at the tip, not extending into a bristle . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. selaginoides

1. Cones sessile, rectangular in cross-section; leaves with grooved midvein, without teeth near the tip,extending into an apical bristle.

2. Leaves distinct from stem in colour, their bases abruptly adnate, not decurrent (except on the topof the branches) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. wallacei

2. Leaves about the same colour as the stem, strongly decurrent.

3. Leaves adnate to the stem for nearly 1⁄2 of their length; plants with long branches, formingloose mats, usually epiphytic on stems and branches of trees . . . . . . . . . . . . . . . . S. oregana

3. Leaves adnate to the stem for less than 1⁄4 of their length; plants with short branches, formingdense mats, terrestrial on soil or rocks.

4. Leaf bristles less than 1 mm long; branches intricate, with scattered roots S. sibirica

4. Leaf bristles at least 1 mm long; branches discrete, without scattered roots S. densa

Selaginella densa Rydb.Compact selaginellaHabitat/Range: Dry soil and rocks in the steppe vegetation zone and from the montane to alpine

zones; frequent in SC BC, infrequent N to 56°N; E to SE MB and S to SD, W TX, NM, AZ and NCA.

Notes: A variable species; two varieties occur in BC.1. Sporophylls denticulate to piliform-ciliate from the base to the apex . . . . . . . . . . var. densa1. Sporophylls ciliate only at the base . . . . . . . . . . . . . . . . . . . . . var. scopulorum (Maxon) Tryon

Selaginella oregana D.C. EatonOregon selaginellaHabitat/Range: On trunks and branches of trees, and moist shady rocks in the lowland zone; rare in

coastal BC, known only from the Queen Charlotte Islands and Broken Islands; S to N CA.

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Selaginellaceae/Adiantaceae

Selaginella selaginoides (L.) LinkMountain-mossHabitat/Range: Wet inundated places, peat bogs or boggy lake shores in the lowland to montane zones;

infrequent throughout BC, occurring mostly on the coast and in the Rocky Mountains; circumpolar, Nto AK, YT and NT, E to NF and S to ID, WY and CO; Eurasia.

Selaginella sibirica (Milde) Hieron.Northern selaginellaHabitat/Range: Dry rocks or rocky soil in the alpine zone; rare in N BC; amphiberingian, N to AK, YT and

W NT; E Asia.

Selaginella wallacei Hieron.Wallace’s selaginellaHabitat/Range: Dry rocks and rocky soil from the lowland and steppe vegetation to subalpine zones;

frequent in S BC south of 52°N; E to SW AB and S to MT and N CA.

PTEROPSIDA

ADIANTACEAE

1. Sori clearly separate, borne on underside of reflexed margins of pinnae; pinnules rhomboic-ovate or fan-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adiantum

1. Sori continuous along the margin of the reflexed frond segment; pinnules more or less oblong or linear-lanceolate.

2. Sori on the lower surface of the frond, not covered by the leaf margin; backs of fronds covered with whiteor yellowish powder . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pentagramma

2. Sori on or near the margin of the lower surface of the frond, covered by inrolled indusium-like margin, orapparent margin; back of fronds without white or yellowish powder.

3. Stipes herbaceous, green except at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cryptogramma

3. Stipes wiry, dark and shining.

4. Blades more or less woolly beneath; segments small, the ultimate segments of the leaves lessthan 5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilanthes

4. Blades glabrous (or hairy, but not woolly in Pellaea atropurpurea); the ultimate segments morethan 5 mm long.

5. Fronds deltoid, not much longer than wide; segments linear, narrowly elongate, upper surfacesstriate, shiny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aspidotis

5. Fronds lanceolate or ovate, considerably longer than broad; segments lanceolate to oblong-ovate, upper surfaces not striate or shiny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pellaea

ADIANTUM

1. Rachis appearing branched into ± equal branches; ultimate pinnules oblong, fanshaped, 2-4 times longerthan wide, attached to straight leaf axes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. aleuticum

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Adiantaceae

1. Rachis unbranched; ultimate pinnules round to broadly oblanceolate, not more than 2 times longer thanwide, attached to conspicuously zig-zag flexed leaf axes . . . . . . . . . . . . . . . . . . . . . . . . A. capillus-veneris

Adiantum aleuticum (Rupr.) Paris (A. pedatum L. ssp. aleuticum [Rupr.] Calder & Taylor, ssp. calderi Cody,var. aleuticum Rupr., and var. subpumilum Wagner in Wagner & Boydston)

Maidenhair or northern maidenhair fernHabitat/Range: Moist forests, rocks, rocky scree, and banks in the lowland and montane zones; frequent

throughout BC, mostly S of 55°N; amphiberingian, N to AK, E to SW AB and S to CA; disjunct E onserpentines in PQ, NF, and VT; E Asia.

Notes: Small plants with small fronds (5 to 15 cm across) and strongly imbricate pinnules were describedas A. pedatum L. var. subpumilum Wagner in Wagner & Boydston. They clearly belong to this species,but no formal nomenclatural transfer has been done yet. This variety or form is rare on coastal bluffsfrom N Vancouver Island S to Olympic Peninsula, WA.

Adiantum capillus-veneris L.Venus-hair or southern maidenhair fernHabitat/Range: Wet rock cliffs in the montane zone; rare in E BC, known only from Fairmont Hot Springs;

cosmopolitan, disjunct S to SD, VA, NC, FL, TX and CA; Europe, S Asia, Africa, S America, Australia.

ASPIDOTIS

Aspidotis densa (Brackenr.) Lellinger (Cryptogramma densa [Brackenr.] Diels in Engl. & Prantl,Cheilanthes densa [Brackenr.] St.John, Cheilanthes siliquosa Maxon)

Indian’s-dream fernHabitat/Range: Rock outcrops and seepages in the lowland, steppe vegetation and montane zones;

infrequent in BC south of 54°N; disjunct E to PQ and S to MT, ID, and CA.

CHEILANTHES

1. Blades and rachis without scales; pinnules tomentose beneath and thinly villous above . . . . . . . C. feei

1. Blades and rachis with scales; pinnules densely tomentose beneath, glabrous above . . . . C. gracillima

Cheilanthes feei T. MooreSlender lip fernHabitat/Range: Dry calcareous (or other basic) rocks in the montane zone; infrequent in S BC east of the

Coast-Cascade Mountains; E to AB and S to WI, IL, AR, TX, NM, AZ, MX and CA.

Cheilanthes gracillima D.C. Eaton in Torr.Lace fernHabitat/Range: Dry rocks and rock crevices in the lowland, steppe vegetation and montane zones; rare in

C and extreme S BC; S to ID, MT, UT and CA.

CRYPTOGRAMMA29

1. Rhizomes slender, creeping; leaves scattered along the rhizome, delicate; sterile leaves ovate to lanceo-late in the outline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. stelleri

1. Rhizomes stout; leaves tufted, relatively coarse; sterile leaves broadly ovate to almost triangular in theoutline base.

29 Contributed by E.R. Alverson & A. Ceska.

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Adiantaceae

2. Mature leaves soft, thin and translucent when dried, deciduous in autumn; dried leaves of a previousyear not persisting on the plant; the upper side of leaves lacking trichomes . . . . . . . . C. cascadensis

2. Mature leaves coriaceous, opaque, evergreen; dried leaves of a previous year persisting on the plant;the upper side of leaves with cylindrical trichomes on veins.

3. Sterile leaves 2-3 times pinnate, coarsely dissected; their ultimate segments elongated, lanceolate oroblanceolate with 6-12 or more teeth or shallow lobes . . . . . . . . . . . . . . . . . . . . . . C. acrostichoides

3. Sterile leaves 2-4 times pinnate, finely dissected; their ultimate segments ovate or obovate, with 2-6deep lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. sitchensis

Cryptogramma acrostichoides R. Br. in Richards. (C. crispa [L.] R. Br. ex Hook. var. acrostichoides [R. Br.in Richards.] C.B. Clarke, ssp. acrostichoides [R. Br. in Richards.] Hult.)

Parsley fern, or mountain-parsleyHabitat/Range: Dry rocks from the lowland to alpine zones; frequent throughout S BC, less frequent

northward; amphiberingian, N to AK, YT and NT, E to ON and PQ and S to MI, MT, WY, CO, NM, UT,NV and CA; E Asia.

Cryptogramma cascadensis AlversonCascade parsley fernHabitat/Range: Dry to mesic rocks and scree slopes in the alpine zone; rare in BC, known only from

Revelstoke and the Cascade Mountains; S to CA.

Cryptogramma sitchensis (Rupr.) T. Moore (C. acrostichoides R. Br. var. sitchensis [Rupr.] C. Christ.)Sitka parsley fernHabitat/Range: Dry to mesic rocks and talus slopes from the lowland to alpine zones; infrequent in NW

BC; N to AK, YT and NT.

Cryptogramma stelleri (Gmel.) PrantlSlender rock-brakeHabitat/Range: Wet crevices in calcareous rocks from the montane to alpine zones and in the steppe

vegetation zone; infrequent throughout BC, mainly E of the Coast-Cascade Mountains; circumpolar, Nto AK, YT and NT, E to AB, disjunct to ON and E to NF and S to NY, NJ, PA, WV, IL, IA, MT, NM, UTand OR; extreme NE Europe, N Asia.

PELLAEA

1. Stipes and rachis sparsely pilose, dull; pinnae long-stalked, the basal ones with stalks 5-15 mm long; sterileand fertile fronds dissimilar, the fertile exceeding the sterile ones . . . . . . . . . . . . . . . . . . . P. atropurpurea

1. Stipes and rachis glabrous or nearly so, shiny; pinnae short-stalked with stalks 0-6 mm long; sterile andfertile fronds alike . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. occidentalis

Pellaea atropurpurea (L.) LinkPurple cliff-brakeHabitat/Range: Dry to mesic limestone rocks in the montane and subalpine zones; rare in SE BC; E to AB,

SK and disjunct to ON and S to NY, MI, WI, MT, SD, WY, FL, NM, AZ, MX and Guatemala.

Pellaea occidentalis (E. Nelson) Rydb. ssp. simplex (Butters) Gastony (P. glabella var. simplex [E.Nelson] Butters and P. suksdorfiana Butters = ssp. simplex; P. pumila Rydb. and P. glabella var.occidentalis [E. Nelson] Butters = ssp. occidentalis)

Simple or smooth cliff-brake

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Adiantaceae/Aspleniaceae

Habitat/Range: Dry to mesic crevices of calcareous rocks in the steppe vegetation and montane zones;infrequent in SC and SE BC, rare in NE; E to AB, S to WA, and disjunct to NM, UT, CO, and AZ.

Notes: In British Columbia, only the ssp. simplex is known. The other subspecies, ssp. occidentalis, canbe expected in SE BC. The two subspecies can be distinguished as follows:1. Rachis reddish-brown to brown-purple; stipes thick and sturdy, old stipes conspicuous; sporangia

with 32 spores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ssp. simplex (Butters) Gastony1. Rachis golden to dark brown; stipes thin and brittle, old stipes inconspicuous; sporangia with 64

spores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ssp. occidentalis

PENTAGRAMMA

Pentagramma triangularis (Kaulf.) Yatskievych, Windham & Wollenweber (Pityrogramma triangularis[Kaulf.] Maxon)

Goldenback fernHabitat/Range: Dry rocky crevices in the lowland zone; infrequent on SE Vancouver Island and adjacent

coast; S to NM, UT, NV, AZ and CA.

ASPLENIACEAE

ASPLENIUM

1. Stipes green or dark green, deciduous; rachis green throughout . . . . . . . . . . . . . . . . . . . . . . . . . . . A. viride

1. Stipes purplish brown, persistent; rachis dark brown or purplish brown, at least in its lower part.

2. Rachis brown or purplish brown almost to the tip, winged with whitish scarious margins (at 10xmagnification); pinnae arranged flat with the plane of the blade . . . . . . . . . . . . . . . . . . A. trichomanes

2. Rachis brown or purplish brown on the lower part, green on the upper third of the frond, lacking thewhitish scarious margins; pinnae arranged obliquely or perpendicular to the plane of the blade (as inVenetian blinds) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. adulterinum

Asplenium adulterinum MildeCorrupt spleenwortHabitat/Range: Dry to mesic walls of limestone fissures in the subalpine zone; rare on N Vancouver

Island, not known elsewhere in N America; Europe.Notes: This species is an allotetraploid which originated from a hybrid of diploid species A. trichomanes x

A. viride. Our plants may belong to A. adulterinum ssp. presolanense Mokry, Rasbach & Reichstein(Mokry et al. 1986).

Asplenium trichomanes L. (A. melanocaulon Willd.)Maidenhair spleenwortHabitat/Range: Dry to mesic rocks in lowland and montane zones; frequent in coastal BC,

infrequent in the interior; circumpolar, N to S AK, E to AB and disjunct in ON to NF and S toGA, LA, TX, NM, AZ and CA; Eurasia.

Notes: A variable species. Two chromosome races (diploid ssp. tr ichomanes and tetraploid ssp.quadrivalens D.E. Meyer) were reported from N America (Moran 1982). The third race, diploidssp. inexpectans Lovis, was described from Europe. According to Mokry et al. (1986) thissubspecies participated in the origin of our A. adulterinum and thus could be expected in BC.Morphological characters distinguishing these races, however, are very minute.

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Aspleniaceae/Azollaceae/Blechnaceae

Asplenium viride Huds. (A. trichomanes-ramosum L.)Green spleenwortHabitat/Range: Mesic to moist crevices in limestone and other basic rocks from the lowland and steppe

vegetation to subalpine zones; infrequent throughout BC; circumpolar, N to AK, YT and NT, E to ABand disjunct in ON to NF and S to VT, MI, WI, SD, MT, CO, UT, NV and OR; Eurasia.

Notes: Some authors advocate the use of Linne’s name A. trichomanes-ramosum, which should berejected as a trinomial.

AZOLLACEAE

AZOLLA

1. Glochidia without (or with one or two) septae; megaspores reticulate in the basal region; rare in SW BC . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. filiculoides

1. Glochidia with many septae; megaspores pitted in the basal region; rare in SC BC . . . . . . A. mexicana

Azolla filiculoides Lam.Large mosquito fernHabitat/Range: Surface of water, sloughs and ditches in the lowland zone; rare in SW BC (lower Fraser

River Valley); introduced from Europe.

Azolla mexicana K.B. PreslMexican mosquito fernHabitat/Range: Surface of water, sloughs and pools in the montane zone; rare in SC BC, known only from

Salmon Arm and Sicamous; S to WI, IL, MO, TX, NM and CA.

BLECHNACEAE

1. Fertile leaves strongly differentiated from sterile leaves; plants 10-60 cm tall . . . . . . . . . . . . . . Blechnum

1. Fertile leaves not differentiated from sterile leaves; plants 60-120 (or more) cm tall . . . . . . Woodwardia

BLECHNUM

Blechnum spicant (L.) Roth (B. doodioides Hook.)Deer fernHabitat/Range: Mesic to wet forests, peatbogs, and open places in the lowland to montane zones;

frequent in coastal BC, infrequent in SC BC; circumpolar, but widely disjunct, N to AK and S to ID, CA;N Africa, Eurasia.

Notes: Our plants are generally more robust than their European counterpart and are referred by someauthors (e.g., Love & Love 1966, 1968) to the amphiberingian ssp. nipponicum (Kunze) Love & Love.

WOODWARDIA

Woodwardia fimbriata J.E. Smith ex ReesGiant chain fernHabitat/Range: Moist forests and seepy coastal cliffs in the lowland zone; rare in BC (SE Vancouver

Island, Lasqueti and Texada Islands); S to CA, disjunct to AZ and NV.

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DENNSTAEDTIACEAE

PTERIDIUM

Pteridium aquilinum (L.) Kuhn (P. aquilinum var. pubescens Underw. & var. lanuginosum [Bong.] Fern. =ssp. lanuginosum; P. aquilinum var. latiusculum [Desv.] Underw. = ssp. latiusculum)

Bracken fernHabitat/Range: Dry to wet forest margins and openings, peatbogs, logged areas and dry meadows from

the lowland and steppe vegetation to subalpine zones; common in BC north to 55°N (ssp. lan-uginosum) or rare in SC BC (ssp. latiusculum); cosmopolitan, E to W AB, disjunct to MB and ON, andS to GA, TX, NM, AZ and CA; Eurasia, Africa, Australia, S America.

Notes: Most BC plants of this species belong to P. aquilinum ssp. lanuginosum. Two subspecies can bedistinguished as follows:1. Leaves broadly triangular, but rarely ternate; pinnules nearly at right angles to the rachis, their

lower surface densely villous or villous-puberulent; the inner indusium ciliate and sometimes alsopubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ssp. lanuginosum (Bong.) Hult.

1. Leaves mostly ternate; pinnules at an oblique angle to the rachis, their lower surface glabrate andshort-pubescent only on the midrib; the inner indusium glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ssp. latiusculum (Desv.) C.N. Page

DRYOPTERIDACEAE

1. Leaves conspicuously dimorphic, fertile and sterile fronds dissimilar . . . . . . . . . . . . . . . . . . . . . Matteuccia

1. Leaves not dimorphic (or slightly dimorphic in Dryopteris cristata), fertile and sterile fronds similar.

2. Indusia present, at least on young fertile fronds, sometimes only as filamentous segments.

3. Indusia splitting into filamentous or ribbon-like segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . Woodsia

3. Indusia membranaceous or scale-like.

4. Indusia attached in the centre, margins of indusia free.

5. Pinnae margins thin and green; indusia kidney-shaped, with a deep sinus in the middle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dryopteris

5. Pinnae margins thick and pale; indusia shield-shaped, without a deep sinus Polystichum

4. Indusia attached along their margins, lanceolate or pouch-like.

6. Indusia lanceolate, attached along their margin; free margins of indusia entire Athyrium

6. Indusia ovate, pouch-like; free margins of indusia serrate or lacerate . . . . . . . . . Cystopteris

2. Indusia absent.

7. Leaves tufted on a short rhizome that is covered by the conspicuous, persistent, densely clusteredpetiole-bases of previous years.

8. Sori elongate; leaves relatively large, 2-8 dm long, forming a vase-like tuft; persistent petiolebases appressed, coarse, flattened, 3 mm wide or more . . . . . . . . . . . . . . . . . . . . . . . . . Athyrium

8. Sori round; leaves smaller, 0.5-3.5 dm long, not forming a vase-like tuft; persistent petiole basesfree standing, more slender and wiry, not more than 2 mm wide . . . . . . . . . . . . . . . . . . Woodsia

7. Leaves scattered or in small tufts, not accompanied by conspicuous, persistent petiole basesof previous years.

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9. Pinnae of the first order alternate; leaves lanceolate or triangular-lanceolate in the outline,longer than broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cystopteris

9. Pinnae of the first order opposite; leaves triangular in the outline, about as long as broad. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gymnocarpium

ATHYRIUM

1. Indusia crescent-shaped, persistent; fronds well-expanded, flat; pinnae not appearing crowded ordirected sharply towards the apices of the fronds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. filix-femina

1. Indusia inconspicuous, withering early; fronds obliquely folded; pinnae crowded and directedsharply towards the apices of the fronds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. distentifolium

Athyrium distentifolium Tausch ex Opiz ssp. americanum (Butters) Hult. (A. alpestre [Hoppe] Opiznon Clairv., A. distentifolium var. americanum [Butters] Boivin, A. americanum [Butters] Maxon)

Alpine lady fernHabitat/Range: Dry to mesic scree slopes and rocks in the subalpine and alpine zones; infrequent

in S BC, rare northward; N to AK, YT and NT, disjunct in PQ and NF, S to WY, CO, NE and CA.

Athyrium filix-femina (L.) Roth ssp. cyclosorum (Rupr.) C. Christ. in Hult. (Athyrium cyclosorumRupr. ,A. filix-femina var. sitchense Rupr. and var. californicum Butters)

Lady fernHabitat/Range: Mesic to wet alluvial forests, swamps and rock outcrops from the lowland and

steppe vegetation to alpine zones; common throughout BC; amphiberingian, N to AK, YT andNT, E to NF and S to PE, OH, WI, IA, and CA; E Asia.

CYSTOPTERIS

1. Rhizomes short; fronds forming clumps; stipes shorter than the leaf blades; leaf blades lanceolate orovate, about 2-3 times longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. fragilis

1. Rhizomes long-creeping; fronds arising singly at the end of the rhizome; stipes longer than the leafblades; leaf blades triangular, about as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. montana

Cystopteris fragilis (L.) Bernh. (C. dickieana R. Sim)Fragile fernHabitat/Range: Dry to moist rocks and forest margins from the lowland and steppe vegetation to

alpine zones; common throughout BC; cosmopolitan, N to AK, YT and NT, E to NF, throughoutthe US; Eurasia, Africa, New Zealand, Australia, S America, Antarctic Islands.

Notes: This variable species as treated here includes both plants with spiny spores (C. fragiliss.str.) and those with rugose spores. Plants with rugose spores were segregated in Europe asC. dickieana. Ceska & Ceska (unpubl.) found that our populations with rugose spores are notuniform in other characters and agree with C. Haufler (cited in Lellinger 1985), that ‘‘some of theAmerican material ... may not be conspecific with material from the type locality of C.dickieana...’’

Cystopteris montana (Lam.) Bernh. (Rhizomatopteris montana [Lam.] Khokhr.)Mountain bladder fernHabitat/Range: Moist to wet calcareous scree slopes, rock outcrops and damp woods in the

montane and subalpine zones; rare in N BC and in the Rocky Mountains; circumpolar, N to AK,YT and NT, E to AB, disjunct in ON, PQ, NF and S to MT and disjunct in CO; Eurasia.

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DRYOPTERIS

1. Blades densely covered by capitate glands; indusia with glandular margins; plants aromatic . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. fragrans

1. Blades and indusia without glands; plants not aromatic.

2. Fronds strongly dimorphic, fertile fronds taller and narrower than the sterile ones; pinnae markedlyreduced towards the base; segments of the lower pinnae rounded in outline, broadly attached andscarcely or not at all contracted at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. cristata

2. Fronds alike; pinnae not reduced towards the base; segments of the lower pinnae oblong in outline,narrowly attached to the costa and contracted at the base.

3. Sori located close to the leaf margin; apical part of pinnules shallowly crenulate or entire; leaves bluegreen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. marginalis

3. Sori located between the leaf margin and the midrib, never directly at the leaf margin; apical part ofpinnules serrate or dentate; leaves dark green or yellow green, without a blue tinge.

4. Laminae three to four times pinnate; leaf blades not narrowed towards their base.

5. Leaf blades triangular in outline, broadest at the base, lines connecting tips of the lower pinnaeclearly divergent; basal pinnules of the lowest pinna alternate, those pointing down more thantwice as long as the pinnules pointing up . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. expansa

5. Leaf blades elongate deltoid in the outline, lines connecting tips of lower pinnae almost parallel;basal pinnules of the lowest pinna approximated and almost opposite, those pointing down lessthan twice as long as the pinnules pointing up . . . . . . . . . . . . . . . . . . . . . . . . . . D. carthusiana

4. Laminae pinnate-pinnatifid to sometimes twice pinnate-pinnatifid; leaf blades narrowed towardstheir base.

6. Pinnae subsessile, oblong-lanceolate; the lower basal pinnules asymmetrical, with semicor-date base, overlying the main rachis; costa with narrowly lanceolate scales . . . . . D. arguta

6. Pinnae clearly stalked, deltoid-lanceolate; basal pinnules symmetrical with cuneate base;costa with hair-like scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. filix-mas

Dryopteris arguta (Kaulf.) WattCoastal wood fernHabitat/Range: Mesic forest margins and rocky coastal cliffs in the lowland zone; rare in SW BC, known

from the Nanoose Hill area, Denman and Hornby Islands; S to CA, disjunct in AZ.

Dryopteris carthusiana (Vil.) H.P. Fuchs (D. spinulosa [Sw.] Watt, D. austriaca auct. non [Jacq.] Woynar, D.dilatata auct. non Hoffm.)

Toothed wood fernHabitat/Range: Wet forests and swamps in the lowland, steppe vegetation and montane zones; infre-

quent throughout BC; circumpolar, N to YT and NT, E to NF and S to SC, AR, NE, MT, ID and WA;Eurasia.

Dryopteris cristata (L.) A. GrayCrested wood fernHabitat/Range: Wet swamps and meadows in the steppe vegetation and montane zones; rare in C and

SE BC; circumpolar, E to NF and S to NC, TN, KS, MT, ID and WA; Europe, W Asia.

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Dryopteris expansa (K.B. Presl) Fraser-Jenkins & Jermy (D. assimilis S. Walker, D. spinulosa [Sw.] Wattssp. assimilis [Walker] Schidlay, D. austriaca auct. non [Jacq.] Woynar, D. dilatata auct. non Hoffm.)

Spiny or spreading wood fernHabitat/Range: Mesic to wet forests, forest margins and scree slopes from the lowland and steppe

vegetation to subalpine zones; common in BC; circumpolar, N to AK, YT and NT, E to NF and S to MI,MN, WY, CO, ID and CA; Eurasia.

Dryopteris filix-mas (L.) SchottMale fernHabitat/Range: Mesic to moist alluvial forests, on scree and in rocky crevices from the lowland to

subalpine zones; frequent in coastal BC except rare on Vancouver Island, infrequent in SC BC; almostcosmopolitan, but widely disjunct, E to NS and S to VT, MI, SD, TX, NM and CA; Eurasia, S America,S Asia, S Africa.

Dryopteris fragrans (L.) SchottFragrant wood fernHabitat/Range: Dry rocks and scree from the lowland to subalpine zone; infrequent in N BC; circumpolar,

N to AK, E to NF and S to ME, MI, WI and MN; N Asia, N Scandinavia.

Dryopteris marginalis (L.) A. GrayMarginal wood fernHabitat/Range: Moist woods in montane zone; rare in SW BC (Meager Creek Hot Springs); disjunct from

E North America, E from WI, ON, NB to NS, S to GA, AR and OK.

GYMNOCARPIUM

1. Rachis with glands, especially conspicuous at the base of the pinnae; lower leaf blade surfaces glandular. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. jessoense

1. Rachis glabrous; upper and lower leaf blade surfaces glabrous.

2. Lamina most commonly bipinnate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. dryopteris

2. Lamina most commonly tripinnate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. disjunctum

Gymnocarpium disjunctum (Rupr.) Ching (G. dryopteris ssp. disjunctum [Rupr.] Sarvela, Dryopteris dis-juncta [Rupr.] Morton)

Western oak fernHabitat/Range: Moist forests in the lowland and montane zones; frequent throughout coastal and SC BC;

amphiberingian, N to AK, E to AB and S to WY, MT, OR, and WA; Kamchatka, Sakhalin.

Gymnocarpium dryopteris (L.) Newman (Phegopteris dryopteris [L.] Fee, Thelypteris dryopteris [L.]Slosson)

Oak fernHabitat/Range: Wet rocky forests, bog margins and rocky ledges in the montane and subalpine zones;

common throughout S BC, less frequent northward; circumpolar, N to AK, YT and NT, E to NF and S toPA and MI; Eurasia.

Gymnocarpium jessoense (Koidz.) Koidz. ssp. parvulum Sarvela (G. continentale [V. Petrov] Pojark., G.robertianum auct. non [Hoffm.] Newm.)

Nahanni oak fernHabitat/Range: Moist, cool, shale talus slopes in the montane and subalpine zones; rare in NE BC;

amphiberingian, N to AK, YT, and NT, E to ON and S to WI and MN; Asia.Notes: A hybrid with G. dryopteris, G. x intermedium Sarvela has not been reported from BC, but can be

expected in NE BC.

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MATTEUCCIA

Matteuccia struthiopteris (L.) Tod.Ostrich fernHabitat/Range: Moist to wet sandy to silty gravel banks of rivers and streams and alluvial forests in

the lowland and montane zones; infrequent (but locally abundant) throughout BC, absent on theQueen Charlotte Islands and Vancouver Island; circumpolar, N to AK, YT and NT, E to NF and Sto MD, VA, OH, IL, MI and SD; Eurasia.

POLYSTICHUM30

1. Fronds pinnate, the pinnae not deeply cleft again, with only shallow teeth along the margin.

2. Lowest pinnae triangular or subtriangular to broadly trowel-shaped, symmetrical; all pinnaespreading-spinulose; stipe length less than one tenth of the frond; spores spiny P. lonchitis

2. Lowest pinnae ovate to lanceolate-falcate, asymetrical; pinnae incurved-spinulose; stipe lengthmore than one-tenth of the frond; spores with folded or bumpy surfaces, without spines.

3. Pinnae arranged in the plane of the frond, distant; stipe and rachis persistently chaffy withscales more than 1 mm wide; indusia ciliate; pinnae cuneate at the base . . . . . . P. munitum

3. Pinnae folded inwards and oriented about 45° to the plane of the frond, imbricate; stipe and rachis oftennaked with scales (if present) less than 1 mm wide; indusia entire; pinnae oblique at the base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. imbricans

1. Fronds bipinnatifid to bipinnate, the pinnae deeply lobed, incised, or again pinnate.

4. Pinnae not at all spinulose nor apiculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. lemmonii

4. Pinnae apiculate to spinulose.

5. Fronds clearly twice pinnate, the pinnules distinct, sessile or petiolate.

6. Fronds with vegetative buds on the rachis; indusia entire . . . . . . . . . . . . . . . . . . . . . . . . . P. kwakiutlii31

6. Fronds without vegetative buds on the rachis; indusia sparsely ciliate on the margins.

7. Pinnules at the base of the pinnae of unequal length, those pointing up conspicuously larger thanthose pointing down . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. setigerum

7. Pinnules at the base of the pinnae about the same length . . . . . . . . . . . . . . . . . . . . . . . . . P. braunii

5. Fronds bipinnatifid, the pinnules adnate to the costa for more than 2 mm, usually with fused bases.

8. Pinnae lacking filiform scales on either surface.

9. Pinnae acute at the apex, especially at the base of the blade, armed with coarse (visible withoutmagnification) spreading teeth; stipe shorter than one-fourth of the frond; rare in SW BC . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. kruckebergii

9. Pinnae obtuse at apex, especially at the base of the blade, armed with fine, incurved teeth; stipe atleast one-fourth of the frond length; rare in SC BC . . . . . . . . . . . . . . . . . . . . . . . . . . . P. scopulinum

8. Pinnae with filiform scales beneath.

10. Fronds with a vegetative bud on the rachis, usually one third of the way down from the tip.

30 Key adapted from Wagner (1979).31 See Excluded species.

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11. Indusia sparsely ciliate on the margin; infralaminar trichomes with contorted projections . . P.andersonii

11. Indusia entire; infralaminar trichomes simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. kwakiutlii32

10. Fronds without a vegetative bud.

12. Pinnae not incised to the costa, the pinnules connate at least one-fourth their length . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. californicum33

12. Pinnae incised to the costa, the pinnules scarcely connate . . . . . . . . . . . . . . . . . P. setigerum

Polystichum andersonii Hopkins (P. braunii ssp. andersonii [Hopkins] Calder & Taylor)Anderson’s holly fernHabitat/Range: Mesic to moist coniferous forests, lava flows and rocky scree in the lowland to subalpine zones;

frequent in coastal NW BC, infrequent elsewhere; N to AK and S to MT and OR.

Polystichum braunii (Spenner) Fee (P. braunii ssp. purshii [Fernald] Calder & Taylor)Braun’s holly fernHabitat/Range: Mesic to wet alluvial forests, coniferous montane forests, scree and lava flows from the lowland to

subalpine zones; infrequent in BC; circumpolar, N to AK, disjunct in ON to NS and S to ME, PA, MI, WI andID; Eurasia.

Polystichum imbricans (D.C. Eaton) D.H. WagnerNarrow-leaved sword fernHabitat/Range: Dry rock outcrops, forest margins and open forests in the lowland zone; infrequent in SW BC; S

to CA.

Polystichum kruckebergii W.H. WagnerKruckeberg’s holly fernHabitat/Range: Dry to mesic ultramafic rock outcrops in the montane and subalpine zones; rare in SW BC,

disjunct to NW BC (Cassiar Mountains); S to MT, ID, UT and CA.

Polystichum lemmonii Underw. (P. mohrioides [Bory] K.B. Presl var. lemmonii [Underw.] Fern.)Lemmon’s holly fernHabitat/Range: Dry to mesic ultramafic rock outcrops in the montane zone; rare in BC, known only from

Okanagan Highland; S to CA.

Polystichum lonchitis (L.) RothMountain or northern holly fernHabitat/Range: Dry to moist limestone or other basic rocks and rocky scree in the montane to alpine zones;

frequent throughout BC; circumpolar, N to AK, E to NS and S to MI, MT, WY, CO, AZ and CA; Eurasia.

Polystichum munitum (Kaulf.) K.B. PreslSword fernHabitat/Range: Moist to mesic forests in the lowland and montane zones; common in SW coastal BC, infrequent

in the interior N to 54°N; N to AK and S to MT, ID and CA, disjunct in SD.

Polystichum scopulinum (D.C. Eaton) MaxonCrag holly fernHabitat/Range: Dry to mesic shady ultramafic rock outcrops in the montane zone; rare in SC BC, known only

from the Tulameen River valley; S to ID, WY, UT, AZ, and CA, disjunct in PQ.

32 See Excluded species.33 Ibid.

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Polystichum setigerum (K.B. Presl) K.B. Presl (P. braunii ssp. alaskense [Maxon] Calder & Taylor)Alaska holly fernHabitat/Range: Moist to mesic shady forests, rock outcrops and lava flows in the lowland and montane

zones; rare in coastal NW BC; N to AK.

WOODSIA

1. Stipes not articulate above base, joints absent, fracturing unevenly and leaving persistent stipe bases ofuneven length; laminae glandular.

2. Pinnae and rachis with articulate hairs; glands on pinnae both sessile and multicellular, elongate . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . W. scopulina

2. Pinnae and rachis without articulate hairs; glands on pinnae only sessile . . . . . . . . . . . . . W. oregana

1. Stipes articulate above the base with slightly thickened, darker joints, fracturing evenly and leavingpersistent stipe bases of similar length; laminae mostly eglandular or with only occasional glands.

3. Rachis and pinnae with very scattered multicellular, white hairs, chaffy scales absent; stipes straw-yellow to yellowish green, dark brown only at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . W. glabella

3. Rachis and lower side of pinnae with scattered brown and whitish hairs and chaffy scales; stipes andrachis reddish brown to dark purple.

4. Stipes and the lower surfaces of pinnae with numerous reddish brown scales; pinnae of the middlepart of the frond 2-2.5 times longer than wide, those at the base with 3-6 deep incisions on each side. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . W. ilvensis

4. Stipes and the lower surfaces of pinnae chaffy with a few, often light scales; pinnae in the middle partof the frond 1-1.5 times as long as wide, those at the base with 1-2 deep incisions on each side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . W. alpina

Woodsia alpina (Bolton) S.F. GrayAlpine cliff fern, or northern woodsiaHabitat/Range: Moist calcareous rock outcrops in the montane and subalpine zones; rare in N BC;

circumpolar, N to AK, YT and NT, E to NF and S to ME, NY, MI and MN; Eurasia.

Woodsia glabella R. Br.Smooth cliff fern or woodsiaHabitat/Range: Moist rocky crevices in the montane and subalpine zones; rare in E BC; circumpolar, N to

AK, YT and NT, E to NF and S to NH, NY, MI and MN; Eurasia.

Woodsia ilvensis (L.) R. Br.Rusty cliff fern or woodsiaHabitat/Range: Dry rock outcrops and scree slopes in the montane to alpine zones; rare in N and E BC;

circumpolar, N to AK, YT and NT, E to NF and S to NC, IL and MN; Eurasia.

Woodsia oregana D.C. EatonWestern cliff fern, or Oregon woodsiaHabitat/Range: Dry to mesic calcareous rock outcrops and talus slopes in the steppe vegetation

and montane zones; infrequent in BC east of Coast-Cascade Mountains, mostly south of 55°N;E to PQ and S to VT, OK, NM, and CA.

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Dryopteridaceae/Hymenophyllaceae/Marsileaceae/Ophioglossaceae

Woodsia scopulina D.C. EatonMountain cliff fern, or Rocky Mountain woodsiaHabitat/Range: Dry to mesic rocky crevices and scree slopes in the lowland and steppe vegetation to

subalpine zones; frequent in W BC and SC BC, mostly south of 55°N; N to AK and YT, E to PQ and Sto SD, WY, CO, NM, AZ and CA, disjunct in VA, TN and AR.

HYMENOPHYLLACEAE

MECODIUM

Mecodium wrightii (van den Bosch) Copeland (Hymenophyllum wrightii van den Bosch)Wright’s filmy fernHabitat/Range: Wet shady rock cliffs, epiphytic on trees in lowland zone; infrequent in coastal BC,

sporophytes known from the Queen Charlotte Islands, male gametophytes known on the coast fromAK to Long Beach, W Vancouver Island; amphiberingian, N to AK; E Asia.

MARSILEACEAE

MARSILEA

Marsilea vestita Hook. & Grev.Hairy water-cloverHabitat/Range: Inundated lake margins in the steppe vegetation and montane zones; rare in SC BC; E to

SK and S to TN, LA, TX, CA, and MX.

OPHIOGLOSSACEAE34,35

1. Veins free; trophophores lobed to pinnate (rarely simple); sporophores branched, with fully exposedsporangia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Botrychium

1. Veins reticulate; trophophores simple and unlobed; sporophores simple with embedded sporangia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ophioglossum

BOTRYCHIUM

1. Trophophore transformed into a second sporophore, so that the leaf has two sporophores but notrophophore . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. paradoxum

1. Trophophore normally developed without sporangia (or rarely with a few sporangia near the base);sporophore present or absent.

2. Sporophores commonly absent; trophophores 2-4x pinnate; leaves mostly more than 18 cm tall.

3. Leaves wintergreen, thick and leathery; sporophores arising at ground level; trophophoresstalked with segments ovate, shallowly crenulate or entire . . . . . . . . . . . . . . . . . . B. multifidum

34 The leaves of this family are unique in having two parts - a sterile photosynthetic blade (trophophore) and spore-bearingappendage (sporophore) that arises from the common stalk.

35 Editors note: Four species of Botrychium (B. ascendens W.H. Wagner, B. hesperium (Maxon & Clausen) W.H. Wagner &Lellinger, B. montanum W.H. Wagner, and B. pedunculosum W.H. Wagner) have recently been described by Wagner andWagner (1981, 1983, 1986). Since these species are known from only a single or a few collections in British Columbia, andsince their taxonomic status is still questionable, we prefer to follow more traditional taxonomic treatments.

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Ophioglossaceae

3. Leaves deciduous, thin and herbaceous; sporophores arising in middle of leaf; trophophoressessile, the segments lanceolate to linear lanceolate, coarsely toothed . . . . . B. virginianum

2. Sporophores always present; trophophores simple (rare) to 1- or occasionally 2-pinnate; leavesmostly less than 18 cm tall.

4. Trophophore attachments strongly variable, those with simple blades usually attached high onthe leaf, those with lobed, non-ternate leaves in the middle of the leaf, and those with ternateblades near the base of the leaf . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. simplex

4. Trophophore attachments relatively uniform, usually attached in the middle to near the top ofthe leaf.

5. Trophophores ternate, segments linear to narrowly lanceolate . . . . . . . . . . . B. lanceolatum

5. Trophophores not ternate, segments mostly broader.

6. Pinnae or ultimate segments with at least a slight midrib, pinnately constructed.

7. Pinnae mostly at right angles or slightly ascending, imbricate with overlapping mar-gins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. pinnatum

7. Pinnae all in one plane, not imbricate, margins distant . . . . . . . . . B. matricariifolium

6. Pinnae or ultimate segments lacking midribs, broadly to narrowly flabellate, dichot-omously constructed.

8. Lower pinnae broadly flabellate, the anterior and basal margins at an angle of wellover 90°; all segments tending to be contiguous or overlapping . . . . . . . . . B. lunaria

8. Lower segments narrowly flabellate, the anterior and basal margins at angles of 90° orless; all segments tending to be well separated or remote . . . . . . . . . B. minganense

Botrychium lanceolatum (Gmel.) Angstr.Lance-leaved moonwortHabitat/Range: Mesic to wet rocky slopes, meadows and woods in the montane to subalpine zones;

infrequent throughout BC; circumpolar, N to AK, E to NF and S to VA, WV, OH, MI, WI, NM, UT,AZ and OR; Eurasia.

Botrychium lunaria (L.) Sw.Common moonwortHabitat/Range: Mesic to moist woods, meadows and heath from the montane to alpine zones;

infrequent throughout BC; circumpolar, N to AK, E to NF and S to ME, VE, NY, MI, MT, CO, AZand CA; Eurasia, Australia, New Zealand, Patagonia.

Botrychium matricariifolium (A. Br. ex Doell) A. Br. ex KochChamomile moonwortHabitat/Range: Dry to moist woods, thickets, and meadows in the montane zone; rare in SC BC;

circumpolar, E to NF and S to VA and ID; Eurasia.

Botrychium minganense Vict. (B. lunaria var. minganense [Vict.] Dole, B. lunaria ssp. minganense[Vict.] Calder & Taylor)

Mingan moonwortHabitat/Range: Mesic to wet woods and meadows, in the montane to alpine zones; infrequent

throughout BC; N to AK, YT and NT, E to NF and S to NY, MI, WI, UT, NE and CA.

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Ophioglossaceae/Polypodiaceae

Botrychium multifidum (Gmel.) Rupr. (Sceptridium multifidum [Gmel.] Tagawa, B. silaifolium Presl,B. occidentale Underwood, B. ternatum [Thunb.] Swartz var. intermedium Eaton)

Leathery grape fernHabitat/Range: Wet meadows, lake margins, peat bogs, river banks and alluvial forests from the lowland

to montane zones; frequent in BC north to 55°N; circumpolar, N to AK, E to NF and S to VA, OH, IA,NM, CO, AZ and CA; Eurasia.

Botrychium paradoxum W.H. WagnerTwo-spiked moonwortHabitat/Range: Mesic to moist subalpine sedge meadows in the subalpine zone; infrequent in SC BC,

known only from Juniper Creek (Ashnola River) and Mt. Kobau; E to SK and S to MT and UT.

Botrychium pinnatum St. John (B. boreale auct. non [Fries] Milde, B. boreale var. obtusilobum [Rupr.]Broun)

Northwestern moonwortHabitat/Range: Mesic to moist streambanks, meadows and heath communities in the montane and

subalpine zones; infrequent throughout BC; N to AK, YT and NT, E to MT and S to NV and OR.

Botrychium simplex E. Hitchc.Least moonwortHabitat/Range: Vernal pools and ephemeral seepages in the lowland and montane zones; rare in BC

(known only from Vancouver Island); circumpolar, disjunct E to ON, PQ and NF and S to NJ, MI, WI,IA, WY, NM, CO, UT, NV and CA; Eurasia.

Botrychium virginianum (L.) Sw. (Botrypus virginianus [L.] Michx.)Rattlesnake fernHabitat/Range: Wet alluvial forests, swamps and riverbanks, in the lowland, steppe vegetation and

montane zones; infrequent throughout BC; circumpolar, N to AK, YT and NT, E to NF and Sthroughout the USA from FL to CA; Eurasia.

OPHIOGLOSSUM

Ophioglossum pusillum Raf. (O. vulgatum auct. non L., O. vulgatum L. var. pseudopodum [Blake] Farw.,O. alaskanum Britton)

Northern adder’s-tongueHabitat/Range: Periodically flooded wet meadows and lake margins, in the lowland and montane zones;

rare in S BC; N to AK, disjunct E to ON, NB and NS, S to VA, OH, IL, MT and WA.

POLYPODIACEAE

POLYPODIUM

1. Leaves leathery; veins anastomosing; rhizomes white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. scouleri

1. Leaves not leathery; veins free; rhizomes green with yellowish or orange tinge.

2. Leaves with a row of hairs along the rachis and on veins of pinnae on the upper side of the leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. glycyrrhiza

2. Leaves with midveins and rachis glabrous or with occasional hairs on the upper side.

3. Sori without (or with only a few) dark, glandular aborted sporangia (paraphyses) P. hesperium

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Polypodiaceae/Thelypteridaceae

3. Sori with numerous dark, glandular aborted sporangia.

4. Leaves yellow-green, with scattered glandular hairs above; the lower part of the rachis usuallywith a few dark scales; rare in NE BC . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. virginianum

4. Leaves greyish-green, without glandular hairs above; dark scales on the lower part of rachisusually absent; infrequent in coastal SW BC and Fraser River Canyon . . . . . . . . P. amorphum

Polypodium amorphum Suksd. (P. montense Lang)Irregular or Pacific polypodyHabitat/Range: Dry to mesic rocks and rock fissures from the lowland to subalpine zones; infrequent in

coastal SW BC and Fraser River Canyon, rare N to 56°N; S to WY, CO, AZ and CA.

Polypodium glycyrrhiza D.C. Eaton (P. vulgare L. var. occidentale Hook., P. occidentale [Hook.] Maxon)Licorice fernHabitat/Range: Dry and seasonally wet rocks, trees, and soil humus in the lowland and montane zones;

common in coastal BC; N to SE AK and S to CA, disjunct in AZ.Notes: In the Barkley Sound area this species hybridizes with P. scouleri.

Polypodium hesperium Maxon (P. vulgare L. var. columbianum Gilbert)Western polypodyHabitat/Range: Dry to mesic rocks from the lowland and steppe vegetation to subalpine zones; infrequent

throughout BC north to 56°N; E to W AB and S to SD, NM, AZ and CA.

Polypodium scouleri Hook. & Grev.Leathery polypodyHabitat/Range: Mesic to moist oceanside rock cliffs and trunks and branches of trees in the salt spray

zone in the lowland zone; frequent on the W coast of Vancouver Island, rare northward to the southerntip of Queen Charlotte Islands; S to CA.

Notes: In the Barkley Sound area this species hybridizes with P. glycyrrhiza.

Polypodium virginianum L. (P. sibiricum Siplivinskij)Virginia or rock polypodyHabitat/Range: Dry to mesic rocks in the montane zone; rare in NE BC; amphiberigian, N to AK, YT and

NT, E to NF and S to GA, AR, SD, MN; E Asia.

THELYPTERIDACEAE

1. Indusia absent; midribs of pinnae grooved above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phegopteris

1. Indusia present, kidney-shaped; midribs of pinnae not grooved above . . . . . . . . . . . . . . . . . . . Thelypteris

PHEGOPTERIS

Phegopteris connectilis (Michx.) Watt (Thelypteris phegopteris [L.] Slosson)Narrow beech or cowboy fernHabitat/Range: Wet swamps, forests and shaded seepy rock cliffs in the lowland to subalpine zones;

infrequent throughout BC; circumpolar, N to AK, YT and NT, E to NF and S to NC, TN, IL, IA, MN andOR; Eurasia.

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Thelypteridaceae

THELYPTERIS

1. Rhizomes short and stout; stipes with numerous scales along their whole length; rachis with dense rustybrown hairs especially on the upper side; ultimate leaf segments 3-4 mm wide . . . . . T. quelpaertensis

1. Rhizomes elongate and slender; stipes with a few scales at the base; rachis sparsely pubescent withinconspicuous hairs; ultimate leaf segments 1.5-2 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . T. nevadensis

Thelypteris quelpaertensis (Christ) Ching (T. limbosperma auct. non [All.] H.P. Fuchs, Oreopteris quelpaer-tensis [Christ] Holub)

Mountain fernHabitat/Range: Wet cliffs in the montane and subalpine zones; rare in coastal BC; amphiberingian, N to

AK and S to WA; E Asia.

Thelypteris nevadensis (D.C. Eaton) CluteNevada marsh fernHabitat/Range: Gravel bars in the lowland zone (moist in summer, flooded in winter), rare in SW BC,

known only from the Sooke River, S to CA.

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Welsh, S.L. 1974. Anderson’s flora of Alaska and adjacent parts of Canada. Brigham Young Univ. Press,Provo. 724 p.

PRIMULACEAE

Beamish, K.I. 1955. Studies in the genus Dodecatheon of Northwestern America. Bull. Torrey Bot. Club 82:357-366.

Cholewa, A.F. and D.M. Henderson. 1984. Primula alcalina (Primulaceae): a new species from Idaho.Brittonia 31: 59-62.

Constance, L. 1938. A revision of the genus Douglasia Lindl. Amer. Midl. Natur. 19: 249-259.

Gleason, H.A. 1952. The New Britton and Brown Illustrated Flora of the Northeastern United States andadjacent Canada. Part 3. New York Bot. Gard.

Kelso, S. 1987. Primula anvilensis (Primulaceae): a new species from Northwestern Alaska. Syst. Bot. 12:9-13.

Ray, J.D., Jr. 1956. The genus Lysimachia in the New World. III. Biol. Monogr. 243-4: 1-160.

Robbins, G.T. 1944. North American species of Androsace. Amer. Midl. Natur. 32: 137-63.

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142

Suttill, T.A. 1987. Biosystematics and reproductive biology of Dodecatheon pulchellum s.l. (Primulaceae).M.Sc. thesis, Univ. of Victoria.

Thompson, H.J. 1953. The biosystematics of Dodecatheon. Contrib. Dudley Herb. 4: 73-154.

Williams, L.O. 1936. Revision of the western primulas. Amer. Midl. Natur. 17: 741-748.

RANUNCULACEAE

Benson, L. 1948. A treatise on the North American Ranunculi. Amer. Midl. Natur. 40:1-264.

. 1954. Supplement to a treatise on the North American Ranunculi. Amer. Midl. Natur. 52:328-369.

. 1955. The Ranunculi of the Alaskan Arctic Coastal Plain and the Brooks Range. Amer. Midl. Natur.53:242-255.

Boivin, B. 1944. American Thalictra and their old world allies. Rhodora 46:337-377, 391-445, 453-487.

. 1948. Two new Thalictra from western Canada. Can. Field- Natur. 62:167-170.

. 1953. Notes on Aquilegia. Amer. Midl. Natur. 50:509-510.

. 1953. Notulae taxonomicae: I. Myosurus minimus Linne (Ranunculaceae). Bull. Soc. Royale Bot.Belg. 85:331,332.

Boraiah, G. and M. Heimburger. 1964. Cytotaxonomic studies on the new world Anemone (sectionEriocephalus) with woody rootstocks. Can. J. Bot. 42:89-922.

Brink, D.E. 1980. Reproduction and variation in Aconitum columbianum, with emphasis on California popula-tions. Amer. J. Bot. 67(3):263-273.

Calder, J.A. and R.L. Taylor. 1963. A new species of Isopyrum endemic to the Queen Charlotte Islands ofBritish Columbia and its relation to other species in the genus. Madro∼no 17:69-76.

Campbell, G.R. 1952. The genus Myosorus L. (Ranunculaceae) in North America. Aliso 2:389-403.

Cook, C.D.K. 1966. A monographic study of Ranunculus subgenus Batrachium (DC.) A. Gray. Mitt. Bot.Staatssamml. Mench. 6:47-237.

Cook, S.A. and M.P. Johnson. 1968. Adaption to heterogeneous environments I. Variation in heterophylly inRanunculus flammula L. Evolution 22:496-516.

den Hartog, C. 1967. Rev. of ‘‘A monographic study of Ranunculus subgenus Batrachium (DC.) Gray’’ byC.D.K. Cook. Acta Bot. Neerl. 16:203-204.

Denton, M.F. 1978. Ranunculus californicus, a new record for the state of Washington. Madro∼no 25:132.

Drew, W.B. 1936. The North American representatives of Ranunculus, section Batrachium. Rhodora 38:1-47.

Duncan, T. 1980. A taxonomic study of the Ranunculus hispidus complex in the Western Hemisphere. Univ.Calif. Publ. Bot. 77.

Ewan, J. 1945. A synopsis of the North American species of Delphinium. Univ. Colo. Stud. Ser. Biol.D2:55-244.

Fisher, F.J.F., J.A. Rowley and C.J. Marchant. 1973. The biogeography of the western snow-patch Ranunculiof North America. Comptes Rendus, Soc. Biogeogr. 438:32-43.

Fisher, F.J.F., A. Warner, and E.M. Reimer. 1979. Anomalous apetaly: localized character displacement inRanunculus eschscholtzii. Can. J. Bot. 57:2097-2106.

Heimburger, M. 1961. A karyotype study of Anemone drummondii and its hybrid with A. multifida. Can. J. Bot.39:497-502.

Kapor, B.M. and A. Love 1970. Chromosomes of Rocky Mountain Ranunculus. Caryologia 23:575-594.

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143

Morris, M.I. 1972. A biosystematic analysis of the Caltha leptosepala (Ranunculaceae) complex in the RockyMountains. I. Chromatography and cytotaxonomy. Brittonia 24:177-188.

Morris, M.I. 1973. A biosystematic analysis of the Caltha leptosepala (Ranunculaceae) complex in the RockyMountains. III. Variability in seed and gross morphological characteristics. Can. J. Bot. 51:2259-2267.

Padmore, P.A. 1957. The varieties of Ranunculus flammula L. and the status of R. scoticus E.S. Marshall andR. reptans L. Watsonia 4:19-27.

Pringle, J.S. 1971. Taxonomy and distribution of Clematis sect. Atragene (Ranunculaceae), in North America.Brittonia 23:361-393.

Scott, P.J. 1974. The systematics of Ranunculus gmelini and R. hyperboreus in North America. Can. J. Bot.52:1713-1722.

Smit, P.G. 1973. A revision of Caltha (Ranunculaceae). Blumea 21:119-150.

Smit, P.G. and W. Punt. 1969. Taxonomy and pollen morphology of the Caltha leptosepala complex. Proc.Royal Neth. Acad. Sci. ser. C. 72:16-27.

Tutin, T.G. 1964. Ranunculaceae. In T.G. Tutin, et al. (eds.), Flora Europaea. Vol. 1. Univ. Cambridge Press,Cambridge.

RHAMNACEAE

Brizicky, G.K. 1964. The genera of Rhamnaceae in the southeastern United States. J. Arnold Arbor.45:439-463.

Johnston, M.C. 1963. A new name in Ceanothus. Leafl. West. Bot. 10:64.

ROSACEAE

Anderson, J.P. 1947. Alaska and Yukon species of Rubus subgenus Cylactis Focke. Bull. Torrey Bot.Club 74:25-56.

Bailey, L.H. 1941-1945. The genus Rubus in North America. Gentes Herb. 5:1-932.

. 1947. Studies in Rubus. 4. Species studies in Rubus. Gentes Herb. 7:193-349.

. 1949. Rubus studies - review and additions. Gentes Herb. 7:479-526.

Bocher, T.W. 1969. Experimental and cytological studies on plant species. XII. Sibbaldia procumbensand S. macrophylla. Sven. Bot. Tidskr. 63:189-200.

Brown, S.W. 1943. The origin and nature of variability in the Pacific coast blackberries (Rubusursinus Cham. and Schlecht. and R. lemurum sp. nov.). Amer. J. Bot. 30:686-697.

Brunsfeld, S.J. and R.D. Johnson. 1990. Cytological, morphological, ecological and phenologicalsupport for specific status of Crataegus suksdorfii (Rosaceae). Madro∼no 37:274-282.

Clausen, J., D.D. Keck, and W.H. Heisey. 1940. Experimental studies of the nature of species. II.Potentilla glandulosa and its allies. III. Potentilla gracilis and its allies. IV. Potentilla drummondiiand Potentilla breweri. Carnegie Inst. Wash. Publ. 520:26-195.

Cole, D. 1956. A revision of the Rosa californica complex. Amer. Midl. Natur. 55:211-224.

Einset, J. 1947. Chromosome studies in Rubus. Gentes Herb. 7:181-192.

Elkington, T.T. 1969. Cytotaxonomic variation in Potentilla fruticosa. New Phytol. 68:151-160.

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144

Erlanson, E. 1934. Experimental data for a revision of the North American wild roses. Bot. Gaz. 96:197-259.

Fassett, T. 1941. Mass collections: Rubus odoratus and R. parviflorus. Ann. Missouri Bot. Gard. 28:299-374.

Fernald, M.L. 1919. Rubus idaeus and some of its variations in North America. Rhodora 21:89-98.

Gajewski, W. 1957. A cytogenetic study on the genus Geum. Pol. Tow. Bot. Monogr. 4:1-416.

Hancock, J.F., Jr. and R.S. Bringhurst. 1979. Ecological differentation in perennial octoploid species of Fragaria. Amer.J. Bot. 66:367-375.

Hess, W.J. 1969. A taxonomic study of Spiraea pyramidata Greene (Rosaceae). Sida 3:298-308.

Hulten, E. 1945. Studies in the Potentilla nivea group. Bot. Not. 1945:127-148.

. 1959. Studies in the genus Dryas. Sven. Bot. Tidskr. 53:507-542.

Jones, G.N. 1935. The Washington species and varieties of Rosa. Madro∼no 3:120-135.

. 1939. A synopsis of the North American species of Sorbus. J. Arnold Arbor. 20:1-126.

. 1946. American species of Amelanchier. Ill. Biol. Monogr. 20:1-126.

Kohli, R. and J.G. Packer. 1976. A contribution to the taxonomy of the Potentilla pensylvanica complex in NorthAmerica. Can. J. Bot. 54:706-719.

Kruschke, E.P. 1965. Contributions to the taxonomy of Crataegus. Milw. Public Mus. Publ. Bot. 3:1-273.

Landon, S.W. 1975. A new name for Osmaronia cerasiformis (Rosaceae). Taxon 24:200.

Lewis, W.H. 958. Minor forms of North American species of Rosa. Rhodora 60:237-243.

. 1959. A monograph of the genus Rosa in North America. I. R. acicularis. Brittonia 11:1-24.

Matfield, B., J.K. Jones and J.R. Ellis. 1970. Natural and experimental hybridization in Potentilla. New Phytol.69:171-206.

Nordborg, G. 1963. Studies in Sanguisorba officinalis L. Bot. Not. 116:267-288.

. 1966. Sanguisorba L., Sarcopterium Spach, and Bencomia Webb and Berth.: Delimitation and subdivisionof the genera. Opera Bot. 11:1-103.

. 1967. The genus Sanguisorba section Poterium. Experimental studies and taxonomy. Opera Bot.16:1-166.

Palmer, E.J. 1925. Synopsis of the North American Crataegi. J. Arnold Arbor. 6:5-128.

Porsild, A.E. 1947. The genus Dryas in North America. Can. Field-Natur. 61:175-192.

Rousi, A. 1964. Biosystematic studies on the species aggregate Potentilla anserina L. Ann. Bot. Fenn. 2:47-112.

Rydberg, P.A. 1918. Rosa. North Am. Fl. 22:483-533.

Sojak, J. 1969. Monemklatorische Anmerkungen zur Gattung Potentilla. Folia Geobot. Phytotax. 4:205-209.

Staudt, G. 1962. Taxonomic studies in the genus Fragaria: Typification of Fragaria species known at the time ofLinnaeus. Can. J. Bot. 40:869-886.

Taylor, R.L. and S. Taylor. 1978. The genus Rosa in British Columbia. Davidsonia 9:30-43.

Taylor, S. 1976. Amelanchier alnifolia (Nuttall) Nuttall. Davidsonia 7:5-12.

Taylor, T.M.C. 1974. The rose family of British Columbia. B.C. Prov. Mus. Handb. 30, Victoria. 223 p.

Uttall, L.J. 1973. The scientific name of the Alaska Spiraea. Bull. Torr. Bot. Club 100:236-237.

. 1974. The varieties of Spiraea betulifolia. Bull. Torrey Bot. Club 101:35-36.

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145

RUBIACEAE

Dempster, L.T. 1976. Galium mexicanum (Rubiaceae) of Central America and Western North America.Madro∼no 23:378-386.

Moore, R.J. 1975. The Galium aparine complex in Canada. Can. J. Bot. 53:877-893.

Puff, C. 1976. The Galium trifidum group (Galium, sect. Aparinoides, Rubiaceae). Can. J. Bot. 54:1911-1925.

SALICACEAE

Argus, G.W. 1965. An endemic subspecies of Salix reticulata L. from the Queen Charlotte Islands, BritishColumbia. Can. J. Bot. 43: 1021-1024.

. 1965. The taxonomy of the Salix glauca complex in North America. Contrib. Gray Herb., HarvardUniv. 196. 142 p.

. 1973. The genus Salix in Alaska and the Yukon. Nat. Mus. Natur. Sci. Publ. Bot. No. 2. Nat. Mus.Can., Ottawa. 279 p.

. 1983. Salix. J.G. Packer (ed.) Flora of Alberta. Second Edition. Univ. Toronto Press, Toronto.

. 1986. Salix raupii, Raup’s willow, new to the flora of Alberta and the Northwest Territories. Can.Field-Natur. 100: 386-388.

. 1986. The Salix lucida Muhl. and S. reticulata L. complexes in North America. Can. J. Bot. 64:541-551.

. 1986. The genus Salix (Salicaceae) in the Southeastern United States. Syst. Bot. Monogr. No. 9.

Argus G.W. and T.C. Brayshaw. 1987. Noteworthy Records, British Columbia Salix tweedyi. Madro∼no 34: 268.

Brayshaw, T.C. 1965. The status of the black cottonwood (Populus trichocarpa Torrey and Gray). Can. Field-Natur. 79: 91-95.

. 1976. Catkin bearing plants of British Columbia. B.C. Prov. Mus. Occas. Pap. Ser. No. 18. 176 p.

Brunsfeld, S.J. and F.D. Johnson. 1985. Field guide to the willows of east-central Idaho. Univ. Idaho, Collegeof Forestry, Wildlife and Range Sciences. Bulletin 39.

Cronquist, A. 1964. Salix. In C.L. Hitchcock et al. (eds.). Vascular Plants of the Pacific Northwest. Part 2. Univ.of Washington Press, Seattle.

Dorn, R.D. 1975. A systematic study of Salix section Cordatae in North America. Can J. Bot. 53: 1491-1522.

Dorn, R.D. 1977. Willows of the Rocky Mountain States. Rhodora 79: 390-429.

Eckenwalder, E. 1977. North American cottonwoods (Populus, Salicaceae) of sections Abaso and Aigeir. J.Arnold Arb. 58: 193-208.

Meikle, R.D. 1984. Willows and poplars of Great Britain and Ireland. Bot. Soc. British Isles Handb. No. 4.

Moss, E.H. 1959. Flora of Alberta. Univ. of Toronto Press, Toronto.

Viereck, L.A. and E.L. Little, Jr. 1972. Alaska trees and shrubs. U.S. Dept. Agric., For. Ser. Agric. Handb. No.410.

SANTALACEAE

Piehl, M.A. 1965. The natural history and taxonomy of Comandra (Santalaceae). Mem. Torrey Bot. Club22:1-97.

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146

SARRACENIACEAE

Cody, W.J. and S.S. Talbot. 1973. The pitcher plant Sarracenia purpurea L. in the northwestern part of itsrange. Can. Field-Natur. 87:318-320.

Krajina, V.J. 1968. Sarraceniaceae, a new family for British Columbia. Syesis 1:121-124.

Wherry, E.T. 1933. The geographic relations of Sarracenia purpurea. Bartonia 15:1-6.

SAXIFRAGACEAE

Bohm, B.A. 1979. Flavonoids of Tolmiea menziesii. Phytochemistry 18: 1079-1080.

Bohm, B.A. and C.K. Wilkins. 1978. Chemosystematic studies in the Saxifragaceae sensu lato. 8. TheFlavonoids of Elmera racemosa (Watson) Rydberg. Brittonia 30: 327-333.

Calder, J.A. 1959. Studies in the Saxifragaceae, II. Saxifraga sect. Trachyphyllum in North America. Brittonia11: 228-249.

. 1960. Studies in the Saxifragaceae, III. Saxifraga odontoloma and lyallii, and North Americansubspecies of S. punctata. Can. J. Bot. 38:410-435.

Calder, J.A. and D.B.O. Savile. 1959. Studies in the Saxifragaceae - I. The Heuchera cylindrica complex inand adjacent to British Columbia. Brittonia 11:49-67.

Ceska, A. 1975. Additions to the adventive flora of Vancouver Island, British Columbia. Can. Field-Natur. 89:451-502.

Clement, C.J.E. 1987. Marsh saxifrage, Saxifraga hirculus, and diapensia, Diapensia lapponica, in northernBritish Columbia. Can. Field-Natur. 101: 443-445.

Elvander, P.E. 1972. Taxonomy of Saxifraga occidentalis and S. marshallii. Can. J. Bot. 50: 2131-2141.

. 1984. The taxonomy of Saxifraga (Saxifragaceae) Section Boraphila Subsection Integrifoliae inwestern North America. Syst. Bot. Mongr. 3: 1-44.

Gornall, R.J. 1985. A monograph of Boykinia, Peltoboykinia, Bolandra, and Suksdorfia (Saxifragaceae). J.Linn. Soc. Bot. 90: 1-70.

Gornall, R.J. and B.A. Bohm. 1980. The use of flavonoids in the taxonomy of Boykinia and allies (Sax-ifragaceae). Can. J. Bot. 58: 1768-1779.

Kern, P. 1966. The genus Tiarella in western North America. Madro∼no 8: 152-159.

Krause, D.L. and K.I. Beamish. 1973. Notes on Saxifraga occidentalis and closely related species.Syesis 6: 104-114.

Packer, J.G. 1963. The taxonomy of some North American species of Chrysosplenium L., sectionAlternifolia Franchet. Can. J. Bot. 41: 85-103.

Perkins, W.E. 1987. Systematics of Saxifraga rufidula and related species from the Columbia RiverGorge to southwestern British Columbia. Ph.D. thesis, Univ. British Columbia, Vancouver.

Randhawa, A.S. 1972. The distribution of Saxifraga ferruginea and the problem of refugia in north-western North America. Can. J. Bot. 50: 79-87.

Randhawa, A.S. and K.I Beamish. 1970. Observations on the morphology, anatomy, classification andreproductive cycle of Saxifraga ferruginea. Can. J. Bot. 48:299-312.

Savile, D.B.O. 1973. Vegetative distinctions in Canadian species of Mitella and Tiarella. Can. Field-Natur. 87:460-462.

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147

Skelly, R.J. 1988. A new species of Saxifraga (Saxifragaceae) from the Olympic Mountains, Washing-ton, and Vancouver Island, British Columbia, Madro∼no 35: 126-131.

Soltis, D.E. 1984. Autopolyploidy in Tolmiea menziesii (Saxifragaceae) Amer. J. Bot. 71: 1171-1174.

. 1984. Karyotypic relationships among Elmera, Heuchera and Tellima (Saxifragaceae). Syst.Bot. 9: 6-11.

Soltis, D.E. and B.A. Bohm. 1986. Flavonoid chemistry of diploid and tetraploid cytotypes of Tolmieamenziesii (Saxifragaceae). Syst. Bot. 11: 20-25.

Soltis, D.E. and L.H. Riesenberg. 1986. Autopolyploidy in Tolmiea menziesii (Saxifragaceae): geneticinsights from enzyme electrophoresis. Amer. J. Bot. 73: 310-318.

Soltis, P.S. and D.E. Soltis. 1986. Anthocyanin content in diploid and tetraploid cytotypes of Tolmieamenziesii (Saxifragaceae). Syst. Bot. 11: 32-34.

Taylor, R.J. 1971. Biosystematics of the genus Tiarella in the Washington Cascades. Northwest Sci. 45:27-37.

. 1965. The genus Lithophragma (Saxifragaceae). Univ. Calif. Publ. Bot. 37: 1-89.

Wells, E. F. 1984. A revision of the genus Heuchera (Saxifragaceae) in eastern North America. Syst.Bot. Monogr. 3:45-121.

SCROPHULARIACEAE

Alex, J.F. 1962. The taxonomy, history and distribution of Linaria dalmatica. Can. J. Bot. 40:295-307.

Boivin, B. 1952. Quelques Veronica du Canada. Natur. Can. 79:173-176.

Callen, E.O. 1940. Part 1. Studies in the genus Euphrasia L. J. Bot. 78:213-218.

. 1952. Part 3. Studies in the genus Euphrasia L. Rhodora 54:145-156.

Cantelon, J.E., E.J. Curtis and W.M. Malcolm. 1963. Studies on Melampyrum lineare. Ecology44:466-474.

Crosswhite, F.S. 1967. Revision of Penstemon section Habroanthus (Scrophulariaceae) I. Conspectus.Amer. Midl. Natur. 77:1-11.

. 1967. Revision of Penstemon section Habroanthus (Scrophulariaceae) II. Series Speciosi.Amer. Midl. Natur. 77:12-27.

. 1967. Revision of Penstemon section Habroanthus (Scrophulariaceae) III. Series Virgati.Amer. Midl. Natur. 77:28-41.

Douglas, D. 1973. Root parasitism in Castilleja rhexifolia. Arct. Alp. Res. 5:145-147.

Fernald, M.L. and K.M. Wiegand. 1915. The genus Euphrasia in North America. Rhodora 17:181-201.

Ganders, F.R. 1966. Occurrence of the genus Euphrasia in the Pacific Northwest. Madro∼no 18:160.

Grant, A.L. 1924. A monograph of the genus Mimulus. Ann. Missouri Bot. Gard. 11:99-389.

Griffiths, A.J.F., G. Krause and F.R. Ganders. 1977. A leaf spot polymorphism in Collinsia grandiflora(Scrophulariaceae). Can. J. Bot. 55:654-661.

Heckard, L.R. 1962. Root parasitism in Castilleja. Bot. Gaz. 124:21-29.

Hiesey, W.M., M.A. Nobs and O. Bjorkmann. 1971. Experimental studies on the nature of species. V.Biosystematics, genetics, and physiological ecology of the Eryanthe section of Mimulus. CarnegieInst. Wash. Publ. 628. 213 p.

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148

Holmgren, N.H. 1971. A taxonomic revision of the Castilleja viscidula group. Mem. New York Bot. Gard.21(4):1-63.

. 1973a. Castilleja. In C.L. Hitchcock and A. Cronquist. Flora of the Pacific Northwest. Univ. Wash.Press, Seattle.

. 1973b. Five new species of Castilleja (Scrophulariaceae) from the Intermountain region. Bull.Torrey Bot. Club 100:83-93.

Hulten, E. 1958. The amphi-Atlantic plants and their phytogeographical connections. Kongl. Svenska Vet-enskapsakad. Handl. Ser. 4,7:1-340.

. 1961. Two Pedicularis species from N.W. America, P. albertae n. sp. and P. sudetica sensu lat.Sven. Bot. Tidskr. 55:193-204.

Keck, D.D. 1927. A revision of the genus Orthocarpus. Proc. Calif. Acad. Sci. IV. 16:517-571.

. 1945. Studies in Penstemon - VIII: a cytotaxonomic account of the section Spermunculus. Amer.Midl. Natur. 33:128-206.

Keck, D.D. and A. Cronquist. 1957. Studies in Penstemon - IX. Notes on northwestern American species.Brittonia 8:247-250.

Pennell, F.W. 1921. Veronica in North and South America. Rhodora 23:1-22, 29-41.

. 1934. Castilleja in Alaska and northwestern Canada. Proc. Acad. Phila. 86:517-540.

Sell, P.D. and P.F. Yeo. 1962. Some new North American eyebrights (Euphrasia). Regnum Veg. 64:202-203.

Straw, R.M. 1966. A redefinition of Penstemon (Scrophulariaceae). Brittonia 18:80-95.

Taylor, T.M.C. 1974. The figwort family (Scrophulariaceae) of British Columbia. B.C. Prov. Mus. Handb. No.33, Victoria. 237 p.

SOLANACEAE

Ceska, A. 1986. An annotated list of rare and uncommon vascular plants of the Victoria area. Victoria Natur.43:1-14.

Goodspeed, T.H. 1945. Studies in Nicotiana III. A taxonomic organization of the genus. Univ. Calif. Publ. Bot.18:335-344.

Stebbins, G.L. Jr. & E.F. Paddock. 1949. The Solanum nigrum complex in Pacific North America.Madro∼no 10:70-81.

Waterfall, U.T. 1958. A taxonomic study of the genus Physalis in North America north of Mexico.Rhodora 60:107-114, 128-142, 152-173.

URTICACEAE

Bassett, I.J., C.W. Crompton and D.W. Woodland. 1974. The family Urticaceae in Canada. Can. J. Bot.52:503-516.

Hermann, F.J. 1946. The perennial species of Urtica in the United States east of the Rocky Mountains.Amer. Midl. Natur. 35:773-778.

VALERIANACEAE

Dempster, L.T. 1958. Dimorphism in the fruits of Plectritis and its taxonomic implications. Brittonia10:14-28.

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149

VERBENACEAE

Lewis, W.H. and R.J. Oliver. 1961. Cytogeography and phylogeny of the North American Verbena. Amer. J. Bot.48:638-643.

Perry, L.M. 1933. A revision of the North American species of Verbena. Ann. Missouri Bot. Gard. 20:239-358.

VIOLACEAE

Baird, V.B. 1941. Wild violets of North America. Univ. Calif. Press, Berkeley. 225 p.

Baker, M.S. 1935. Studies in western violets. I. Madro∼no 3: 51-57.

. 1936. Studies in western violets. II. Madro∼no 3: 321-239.

. 1940. Studies in western violets. III. Madro∼no 5: 218-231.

. 1949. Studies in western violets. VI. Madro∼no 10: 110-128.

. 1953. Studies in western violets. VII. Madro∼no 12: 8-18.

. 1957. Studies in western violets. VIII. The Nuttalianae continued. Brittonia 9:217-230.

Brainerd, E. 1905. Notes on New England violets - III. Rhodora 7: 245-248.

. 1921. Violets of North America. Vermont Agric. Exp. Sta. Bull. No. 224. Burlington, Vermont.

Clausen, J. 1964. Cytotaxonomy and distributional ecology of western North American violets. Madro∼no 17: 173-197.

Fabijan, D.M., J.G. Packer and K.E. Denford. 1987. The taxonomy of the Viola nuttallii complex. Can. J. Bot. 65:2562-2580.

Harms, V.L., D.F. Hooper and L. Baker. 1985. Four violets new to the Saskatchewan flora and other rare violets ofeast-central Saskatchewan. Can. Field-Natur. 99: 498-502.

McPherson, G.D. and J.G. Packer. 1974. A contribution to the taxonomy of Viola adunca. Can. J. Bot. 52: 985-902.

Russell, N.H. 1955. The taxonomy of the North American acaulescent white violets. Amer. Midl. Natur. 54: 481-494.

Russell, N.H. 1956. Regional variation patterns in the stemless white violets. Amer. Midl. Naturalist 56: 491-503.

Russell, N.H. and F.S. Crosswhite. 1963. An analysis of variation in Viola nephrophylla. Madro∼no 17: 56-65.

Sorsa, M. 1968. Cytological and evolutionary studies in Palustres violets. Madro∼no 19: 165-179.

PTERIDOPHYTA

ADIANTACEAE

Alverson, E.R. 1989. Cryptogramma cascadensis, a new parsley-fern from western North America. Amer. Fern J. 79:95-102.

Brunton, D.F. 1979. Taxonomy, distribution, and ecology of the cliff-brake ferns (Pellaea: Polypodiaceae) in Alberta.Can. Field-Natur. 93:288-295.

Brunton, D.F. 1986. Status of the southern maidenhair fern, Adiantum capillus-veneris (Adiantaceae) in Canada. Can.Field-Natur. 100:404-408.

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150

Cody, W.J. 1983. Adiantum pedatum ssp. calderi, a new subspecies in northeastern North America.Rhodora 85: 93-96.

Eastham, J.W. 1949. Adiantum capillus-veneris in British Columbia. Can. Field-Nat. 63:112-114.

Gastony, G.J. 1988. The Pellaea glabella complex: electrophoretic evidence for the derivations of theagamosporous taxa and revised taxonomy. Amer. Fern. J. 78: 44-67.

Knobloch, I.W. 1976. Morphological characters in Cheilanthes together with a key to North and CentralAmerican species. Flora 165:507-522.

. 1976. Pteridophyte hybrids. Publ. Mus. Mich. State Univ., Biol. Ser. 5(4):273-352.

Lellinger, D.B. 1968. A note on Aspidotis. Amer. Fern J. 58: 140-141.

Paris, C.A. and M.D. Windham. 1988. A biosystematic investigation of the Adiantum pedatum complexin eastern North America. Syst. Bot. 13: 240-255.

Rigby, S.J. and D.M. Britton. 1970. The distribution of Pellaea in Canada. Can. Field-Natur. 84:137-144.

Smith, A.R. 1975. The California species of Aspidotis. Madro∼no 23: 15-24.

Smith, D.M., S.P. Craig, and J. Santarosa. 1971. Cytological and chemical variation in Pityrogrammatriangularis. Amer. J. Bot. 58: 292-299.

Tryon, A.F. 1957. A revision of the fern genus Pellaea section Pellaea. Ann. Missouri Bot. Gard. 44:125-193.

. 1972. Spores, chromosomes and relations of the fern Pellaea atropurpurea. Rhodora 74:220-241.

Tryon, A.F. and D.M. Britton. 1958. Cytotaxonomic studies of the fern genus Pellaea. Evolution 12:137-145.

Wagner, W.H., Jr. and K.E. Boydston. 1978. A dwarf coastal variety of maidenhair fern, Adiantumpedatum. Can. J. Bot. 56: 1726-1729.

Weatherby, C.A. 1920. Varieties of Pityrogramma triangularis. Rhodora 22: 113-120.

Yatskievych, G., M.D. Windham & E. Wollenweber. 1990. A reconsideration of the genus PityrogrammaLink (Adiantaceae) in the southwestern United States. Amer. Fern J. 80: 9-17.

ASPLENIACEAE

Lovis, J.D. 1964. The taxonomy of Asplenium trichomanes in Europe. Fern Gaz. 9:147-160.

Mokry, F., H. Rasbach and T. Reichstein. 1986. Asplenium adulterinum Milde subsp. presolanensesubsp. nova (Aspleniaceae, Pteridophyta). Bot. Helvetica 96: 7-18.

Moran, R.C. 1982. The Asplenium trichomanes complex in the United States and adjacent Canada.Amer. Fern J. 72: 5-11.

AZOLLACEAE

Brunton, D.F. 1986. Status of the mosquito fern, Azolla mexicana Salviniaceae, in Canada. Canad.Field-Natur. 100: 404-408.

Svenson, H.K. 1944. The new world species of Azolla. Amer. Fern J. 34: 69-84.

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151

BLECHNACEAE

Cody, W.J. 1963. Woodwardia in Canada. Amer. Fern J. 53: 17-27.

Love, A. and D. Love. 1966. The variation of Blechnum spicant. Sven. Bot. Tidskr. 62: 186-196.

. 1968. Cytotaxonomy of Blechnum spicant. Collect. Bot. 7: 665-676.

DENNSTAEDTIACEAE

Cody, W.J. and C.W. Crompton. 1975. The biology of Canadian weeds: 15. Pteridium aquilinum (L.) Kuhn.Can. J. Plant Sci. 55: 1059-1072.

Tryon, R.M. 1941. A revision of the genus Pteridium. Rhodora 43: 1-31, 37-67.

DRYOPTERIDACEAE

Blasdell, R.F. 1963. A monographic study of the fern genus Cystopteris. Mem. Torrey Bot. Club 21: 1-102.

Boivin, B. 1966. Etudes pteridologiques. III. Variations du Woodsia oregana. Bull. Soc. Bot. France 113:407-409.

Britton, D.M. 1962. Dryopteris dilatata (Hoffm.) A. Gray in North America. Rhodora 64: 207-212.

. 1972. Spinulose wood ferns in western North America. Can. Field-Natur. 86: 241-247.

. 1972. Spore ornamentation in the Dryopteris spinulosa complex. Can. J. Bot. 50: 1617-1621.

. 1977. The fern Woodsia obtusa (Spreng.) Torrey in Ontario. Can. Field-Natur. 91: 84-85.

Britton, D.M. and A.C. Jermy. 1974. The spores of Dryopteris filix-mas and related taxa in North America. Can.J. Bot. 52: 1923-1926.

Brown, D.F.M. 1964. A monographic study of the fern genus Woodsia. Nova Hedwigia 16: 1-154.

Carlson, T.M. and W.H. Wagner. 1982. The North American distribution of the genus Dryopteris. Contrib. Univ.Mich. Herb. 15: 141-162.

Cody, W.J. and D.M. Britton. 1984. Polystichum lemmonii, a rock shield-fern new to British Columbia andCanada. Can. Field-Natur. 98: 375.

Crane, F.W. 1960. A key to American Dryopteris based on characters of the perispore. Amer. Fern J.50: 270-275.

Fuchs, H.P. 1974. The correct name of the alpine lady fern. Candollea 29: 181-205; correction,Candollea 29: 249.

Hagenah, D.J. 1961. Spore studies in the genus Cystopteris. I. The distribution of Cystopteris with non-spiny spores in North America. Rhodora 63: 181-193.

Jermy, A.C. and L. Harper. 1971. Spore morphology of the Cystopteris fragilis complex. Brit. FernGazette 10: 211-213.

Maxon, W.R. 1918. Polystichum andersoni and related species. Amer. Fern J. 8: 33-37.

Prange, R.K. and P. von Aderkas. 1985. The biological flora of Canada 6. Matteuccia struthiopteris (L.)Todaro, ostrich fern. Can. Field-Natur. 99: 517-532.

Pryer, K.M. and D.M. Britton. 1983. Spore studies in the genus Gymnocarpium. Can. J. Bot. 61: 377-388.

Sarvela, J. 1978. A synopsis of the fern genus Gymnocarpium. Ann. Bot. Fenn. 15: 101-106.

. 1980. Gymnocarpium hybrids from Canada and Alaska. Ann. Bot. Fenn. 17: 292-295.

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152

Sarvela, J., D.M. Britton and K. Pryer. 1981. Studies on the Gymnocarpium robertianum complex in North America.Rhodora 83: 421-431.

Sorsa, P. 1980. Spore morphology of the fern genus Gymnocarpium and its relations to the taxonomy. Ann. Bot. Fenn.17: 86-90.

von Aderkas, P. 1983. Studies of gametophytes of Matteuccia struthiopteris (ostrich fern) in nature and in culture. Can.J. Bot. 61: 3267-3270.

. 1984. Economic history of ostrich fern, Matteuccia struthiopteris, the edible fiddlehead. Econ. Bot. 38:14-23.

Wagner, D.H. 1979. Systematics of Polystichum in western North America north of Mexico. Pteridologia 1: 1-64.

Wagner, W.H., Jr. 1973. Reticulation of holly ferns (Polystichum) in the western United States and adjacent Canada.Amer. Fern J. 63: 99-115.

Widen, C.J. and D.M. Britton. 1971. A chromatographic and cytological study of Dryopteris dilatata in North Americanand eastern Asia. Can. J. Bot. 49: 247-258.

. 1971. A chromatographic and cytological study of Dryopteris filix-mas and related taxa in North America.Can. J. Bot. 49: 1589-1600.

EQUISETACEAE

Cody, W.J. and V. Wagner. 1981. The biology of Canadian weeds. 49: Equisetum arvense L. Can. J. Plant Sci. 61:123-133.

Hauke, R.L. 1960. The smooth scouring rush and its complexities. Amer. Fern J. 50: 185-193.

. 1963. A taxonomic monograph of the genus Equisetum subgenus Hippochaeta. Beih. Nova Hedwigia 8:1-123.

. 1967. A systematic study of Equisetum arvense. Beih. Nova Hedwigia 13: 81-109.

. 1974. The taxonomy of Equisetum: an overview. New Bot. 1: 89-95.

. 1978. A taxonomic monograph of Equisetum subgenus Equisetum. Nova Hedwigia 30: 385-455.

Page, C.N. 1972. An assessment of interspecific relationships in Equisetum subgenus Equisetum. New Phytol.71:355-369.

HYMENOPHYLLACEAE

Cortes, L.D. and V.J. Krajina. 1968. Mecodium wrightii on Vancouver Island. Amer. Fern J. 58: 181.

Iwatsuki, I. 1961. The occurrence of Mecodium wrightii in Canada. Amer. Fern J. 51: 141-144.

Taylor, T.M.C. 1967. Mecodium wrightii in British Columbia and Alaska. Amer. Fern J. 57: 1-6.

ISOETACEAE

Boivin, B. 1961. Isoetes echinospora Durieu in North America. Amer. Fern J. 51:83-85.

Dorn, R.D. 1972. The nomenclature of Isoetes echinospora and Isoetes muricata. Amer. Fern J. 62:80-81.

Fuchs, H.P. 1962. Nomenklatur, Taxonomie und Systematik der Gattung Isoetes Linnaeus in geschichtlicherBetrachtung. Beih. Nova Hedwigia 3: 1-103.

Pfeiffer, Norma E. 1922. Monograph of the Isoetaceae. Ann. Missouri Bot. Gard. 9: 79-232.

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153

LYCOPODIACEAE

Beitel, J.M. 1979. Clubmosses (Lycopodium) in North America. Fiddlehead Forum 6: 1-8.

Gillespie, J.P. 1962. A theory of relationships in the Lycopodium inundatum complex. Amer. Fern J.52:19-26.

Hickey, R.J. 1977. The Lycopodium obscurum complex in North America. Amer. Fern J. 67: 45-48.

Holub, J. 1964. Lycopodiella, novy rod radu Lycopodiales. Preslia 36:16-22.

. Diphasiastrum, a new genus in Lycopodiaceae. Preslia 47:97-110.

Love, A., and D. Love. 1958. Cytotaxonomy and classification of the Lycopods. Nucleus 1:1-10.

Wilce, J.H. 1965. Section Complanata of the genus Lycopodium. Beih. Nova Hedwigia 19: 1-233.

MARSILEACEAE

Johnson, D.M. 1986. Systematics of the New World species of Marsilea (Marsileaceae). Syst. Bot.Monogr. 11: 1-87.

OPHIOGLOSSACEAE

Clausen, R.T. 1938. A monograph of the Ophioglossaceae. Mem. Torrey Bot. Club 19: 1-177.

Stevenson, D.W. 1975. Taxonomic and morphological observations on Botrychium multifidum(Ophioglossaceae). Madro∼no 23:198-204.

Wagner, W.H., Jr. and L.P. Lord. 1956. The morphological and cytological distinctness of Botrychiumminganense and B. lunaria in Michigan. Bull. Torrey Bot. Club 83: 261-281.

Wagner, W.H., Jr. and F.S. Wagner. 1981. New species of moonworts, Botrychium subg. Botrychium(Ophioglossaceae), from North America. Amer. Fern J. 71: 20-30.

. 1983. Two moonworts of the Rocky Mountains: Botrychium hesperium and a new speciesformerly confused with it. Amer. Fern J. 73: 53-62.

. 1986. Three new species of moonworts (Botrychium subgen. Botrychium) endemic inwestern North America. Amer. Fern J. 76:33-47.

POLYPODIACEAE

Lang, F.A. 1971. The Polypodium vulgare complex in the Pacific Northwest. Madro∼no 21: 235-254.

SELAGINELLACEAE

Tryon, R.M. Jr. 1955. Selaginella rupestris and its allies. Ann. Missouri Bot. Gard. 42: 1-99.

THELYPTERIDACEAE

Morton, C.V. 1963. The classification of Thelypteris. Amer. Fern J. 53:149-154.

Mulligan, G.A. and W.J. Cody. 1979. Chromosome numbers in Canadian Phegopteris. Can. J. Bot. 57:1815-1819.

Smith, A.R. 1971. Chromosome numbers of some New World species of Thelypteris. Brittonia 23:354-360.

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APPENDIX

EXCLUDED SPECIES

DICOTYLEDONS

PRIMULACEAE

Primula veris L. This species was reported by Taylor & MacBryde (1977) and Scoggan (1979) but no recentspecimens have been seen in the herbaria examined. It is probably a garden escape which does notpersist.

RANUNCULACEAE

Aconitum compactum (Reichenbach) Gayer. A garden escape, cited by Brayshaw (1989), which does notpersist in our area.

Anemone deltoidea Hook. Collections cited by Macoun (1883-1890) and Raup (1934) have been reidentified(Scoggan 1978).

Anemone virginiana L. var. cylindroidea Boivin. Cited by Taylor and MacBryde (1977) but not part of ourflora.

Aquilegia vulgaris L. This common garden plant does not persist outside of cultivation in our area.

Clematis alpina (L.) P. Mill. Reported by Boivin (1966-1967) and Taylor and MacBryde (1977), this introduc-tion has been collected only once and is probably not persistent.

Clematis hirsutissima Pursh (C. douglasii Hook.). Cited as occurring in our range by Abrams (1944) andHitchcock et al. (1964) but yet to be recorded in BC.

Coptis occidentalis (Nutt.) T.& G. This species, reported by Hitchcock et al. (1964) and subsequently treatedby Taylor and MacBryde (1977) and Brayshaw (1989), does not reach our range.

Delphinium nuttallii A. Gray. Erroneously reported by Henry (1915) and Abrams (1944).

Nigella damascena L. This ornamental garden plant, cited by Brayshaw (1989), rarely escapes in our area.

Paeonia brownii Dougl. Cited by Macoun (1883-1890) and Henry (1915) but not part of our flora.

Ranunculus aquatilis L. var. aquatilis. Although Brayshaw (1989) cites this European taxon as occurring inBC, there is not adequate research justifying such a treatment.

Ranunculus bulbosus L. (R. tuberosus Horn.). Cited by Scoggan (1978) and Brayshaw (1989), this specieshas not been collected since 1890.

Ranunculus hebecarpus H.& A. Reported from ballast at Nanaimo by Macoun (1883-1890) but neverrecollected.

Ranunculus micranthus (A. Gray) Nutt. ex T.& G. The collections reported by Macoun (1883-1890) arebased upon specimens of R. abortivus (Scoggan 1978).

ROSACEAE

Alchemilla xanthochlora Rothman. Cited in Taylor (1973), but the specimen, in UBC, upon which this wasbased was re-identified as A. subcrenata.

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Cotoneaster franchetii Boiss. Cited in Taylor & MacBryde (1977), but no specimens seen.

Cotoneaster microphyllus Wall. ex Lindl. Cited in Taylor & MacBryde (1977), but no specimens seen.

Crataegus laevigata (Poir.) DC. Cited in Taylor & MacBryde (1977), but specimens are referrable toC. monogyna.

Filipendula rubra (Hill) B.L. Robins. Cited in Taylor & MacBryde (1977), but no specimens seen.

Potentilla arguta Pursh ssp. arguta. Cited in Taylor and MacBryde (1977), but no specimens seen.

Potentilla multifida L. Cited in Taylor (1973) and Taylor & MacBryde (1977), but no specimens seen.

Rosa woodsii Lindl. ssp. woodsii. Cited in Taylor and MacBryde (1977), this prairie subspecies is notknown from BC.

Rubus bifrons Vest ex Trautv. Cited in Taylor and MacBryde (1977), but no specimens seen. Therecord is likely based on misidentifications of Rubus allegheniensis.

Sanguisorba annua (Nutt. ex Hook.) T.& G. A specimen cited from BC in Boivin (1966) is S.occidentalis according to Scoggan (1978).

RUBIACEAE

Asperula arvensis L. Cited by Taylor and MacBryde (1977) from an old collection at Essondale. Thisspecies is not established in BC.

Galium palustre L. Reported for BC by Taylor and MacBryde (1977), but no material seen.

SALICACEAE

Populus alba L. Cultivated in British Columbia but not naturalized.

Populus nigra L. var. italica DuRoi. Cultivated in British Columbia but not naturalized.

Salix cordata Muhl. non Michx. This name, a synonym of the eastern American S. eriocephala Michx.,was used by Henry (1915). He was probably referring to S. prolixa.

Salix fluviatilis Nutt. This species, reported by Henry (1915), is unconfirmed for BC. Its taxonomicstatus is also uncertain.

Salix lasiolepis Benth. Reported for Hazelton by Cronquist (1964) and repeated by Taylor andMacBryde (1977). No specimens from BC have been located.

SANTALACEAE

Comandra umbellata (L.) Nutt. var. umbellata. This more eastern variant was cited by Hitchcock etal. (1964) from BC but most authors treat our northern material as var. pallida.

SAXIFRAGACEAE

Chrysosplenium glechomaefolium Nutt. ex T.& G. This species of the US west coast, first reportedby Henry (1915) and taken up by Hitchcock et al. (1961) and Taylor and MacBryde (1977), doesnot occur in BC.

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Saxifraga aizoon Jacq. Cited by Scoggan (1978) but no BC material available.

Saxifraga davurica Willd. ssp. grandipetala (Engl. & Irmsch.) Hult. Mapped by Hulten (1968) for BC butnot documented.

Saxifraga foliolosa R. Br. in Parry. Reported by Taylor and MacBryde (1977) but no material seen.

Saxifraga radiata Small. This species, reported by Taylor and MacBryde (1977), is not part of our flora.

Saxifraga rhomboidea Greene. Cited for BC by Hitchcock et al. (1961) and Scoggan (1978) but not occurringin our range.

SCROPHULARIACEAE

Antirrhinum majus L. Cited by Taylor and MacBryde (1977), this garden escape is not established in ourregion.

Besseya rubra (Dougl. ex Hook.) Rydb. The report of this species from SE BC (Henry 1915) was probablybased upon B. wyomingensis.

Castilleja flava S. Wats. Cited by Scoggan (1979) but no good BC material seen of this US species.

Digitalis ambigua Murr. (D. grandiflora P. Mill.). This garden escape, reported for BC by Scoggan (1979) isnot established in our area. It is known from a single 1954 collection at Aldergrove.

Digitalis lanata Ehrh. Scoggan (1979) cited a single 1954 collection from Sooke. This garden escape doesnot persist in our region.

Euphrasia mollis (Ledeb.) Wettst. The report of this species in N BC (Rydberg 1922) has not beensubstantiated.

Kickxia spuria (L.) Dumort. Cited by Eastham (1947) and Scoggan (1979) but known from only asingle collection at Duncan.

Limosella subulata Ives. Reports of this species in Taylor (1974) and Taylor and MacBryde (1977), arebased on misidentifications of L. aquatica (Straley et al. 1985).

Linaria macroccana Hook. f. Cited by Taylor and MacBryde (1977) but no BC material seen.

Linaria pinifolia (Poir.) Thell. (L. reticulata [Sm.] Desf.). The only report of this species from BC (Grohand Frankton 1949) is now over 40 years old and no recent material is available.

Lindernia dubia (L.) Pennell. Reported for BC by Scoggan (1979) but no substantiating material available.

Mimulus suksdorfii A. Gray. This species of the W US, reported for BC by Rydberg (1922), does not extend into ourregion.

Misopates orontium (L.) Raf. (Antirrhinum orontium L.). Cited by Taylor and MacBryde (1977) but no material seen.

Orthocarpus erianthus Benth. This introduction, collected by J. Macoun in 1908 at Victoria (Scoggan 1979), hasnever been recollected.

Orthocarpus purpureus Benth. Cited by Eastham (1947) from the Victoria area but neither this collection nor anyother seen.

Penstemon attenuatus Dougl. ex Lindl. Cited in Taylor (1974) and Taylor & MacBryde (1977), but probably basedon misidentifications of P. albertinus.

Penstemon deustus Dougl. ex Lindl. Cited in Henry (1915), but no specimens known.

Penstemon montanus Greene. Cited in Straley et al. (1985), but probably based on misidentifications of P. ellipticus.

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Penstemon triphyllus Dougl. ex Lindl. Cited in Rydberg (1922), but no specimens known.

Penstemon venustus Dougl. ex Lindl. Cited in Macoun (1884), but this was based on P. lyallii.

Veronica agrestis L. Cited in Henry (1915), but no recent records known.

SIMAROUBACEAE

Ailanthus altissima (P. Mill.) Swingle. This horticultural species, cited by Taylor and MacBryde (1977), doesnot persist in our region.

SOLANACEAE

Lycopersicon esculentum P. Mill. Cited in Taylor & MacBryde (1977) and Scoggan (1979) and althoughfound in garbage dumps and waste places, not truly established.

Nicandra physalodes (L.) J. Gaertn. Cited in Taylor & MacBryde (1977), based on a single plant (specimenin V) from a garden dump and not known to have become established or to have been found elsewhere.

Nicotiana attenuata Torr. ex S. Wats. Cited in Taylor & MacBryde (1977) and Scoggan (1979), based on oldrecords from Spences Bridge, but no collections known since 1923.

Physalis pubescens L. Cited in Scoggan (1979), based on an 1884 record from Cache Creek, but there areno recent records known.

Salpichroa rhomboidea (Gillis & Hook.) Miers. Ceska (1986) cited this species from a 1916 Victoriacollection but no recent material known.

Solanum tuberosum L. Cited in Scoggan (1979), based on a record in CAN from Liard Hot Springs, but notthought to ever become established.

TAMARICACEAE

Tamarix parviflora DC. Reported by Taylor and MacBryde (1977) but not persisting in our region.

VIOLACEAE

Viola macloskeyi Lloyd. This species of OR and CA, cited by many authors (e.g., Hitchcock and Cronquist1973, Taylor and MacBryde 1977, Scoggan 1978) does not occur in BC.

Viola nuttallii Pursh. This species, reported by Taylor and MacBryde (1977), occurs only east of thecontinental divide.

Viola simulata Baker. Described by Baker from material collected at Shawnigan Lake (Henry 1915) andplaced in synonymy with V. langsdorfii by Scoggan (1978). Prior to Baker’s description of V. simulata thetype material had been determined as V. langsdorfii. The status of the species remains in doubt.

VITACEAE

Parthenocissus inserta (Kern.) Fritsch. This horticultural species is known from only a singlecollection at Oliver.

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PTERIDOPHYTA

ASPLENIACEAE

Phyllitis scolopendrium (L.) Newm. (Asplenium scolopendrium L.). Collected in Port Eliza Inlet, Wcoast of Vancouver Island by S.O. Charcott in 1980. The cytology of this population indicates thatthis is an introduction of a European taxon (Cody & Britton 1989).

DRYOPTERIDACEAE

Polystichum californicum (D.C. Eaton) Diels. Collected on Texada Island by W.B. Anderson in 1937.No recent collections.

Polystichum kwakiutlii D.H. Wagner Recently described species based on a specimen collected inAlice Arm by A.D. York in 1935. No recent collections.